Cardamine - Brassibase

CARDAMINE Linnaeus, Sp. Pl. 2: 654. 1753.
Ihsan A. Al-Shehbaz and Karol Marhold
Tribe: Cardamineae Dumort., Fl. Belg.: 124. 1827.
Name derivation: Greek kardamon, used by Dioscorides for some species of Brassicaceae.
Lectotype species (designated by Britton & Brown, Ill. Fl. N.U.S., ed. 2, 2: 183. 1913): C.
pratensis L.
Dentaria Linnaeus, Sp. Pl. 2: 653. 1753. Lectotype species (designated by Britton & Brown, Ill.
Fl. N.U.S., ed. 2, 2: 187. 1913: D. pentaphyllos L.
Dracamine Nieuwland, Amer. Midl. Naturalist 4: 40. 1915. Type species: not designated.
Heterocarpus Philippi, Bot. Zeitung 14: 641. 1856, not Wight (1853). Type species: H.
fernandezianus R. A. Philippi.
Ghinia Bubani, Fl. Pyrenaea 3: 158. 1901, non Schreber (1789). Type species: not designated.
Iti Garnock-Jones & P. N. Johnson, New Zealand J. Bot. 25: 603. 1988. Type species: I. lacustris
Garnock-Jones & P. N. Johnson.
Loxostemon J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5: 129. 1861. Type species: L.
pulchellus J. D. Hooker & Thomson.
Porphyrocodon W. J. Hooker in Bentham & J. D. Hooker, Gen. Pl. 1: 79. 1862. Type species: P.
pictus (W. J. Hooker) B. D. Jackson.
Pteroneurum de Candolle, Mém. Mus. Hist. Nat. 7: 231. 1821. Type species: not designated.
Sphaerotorrhiza (O. E. Schulz) Khokhrjakov, Fl. Magadansk. Obl. 235. 1985. Type species: S.
trifida (Poiret ex Lamarck) Khokhrjakov.
Herbs, annual, biennial, or perennial, rhizomatous or tuberous, sometimes with axillary
bulbils. Trichomes absent or simple. Multicellular glands absent. Stems erect or prostrate,
leafy or rarely leafless and plant scapose. Basal leaves petiolate, rosulate or not, simple and
entire, toothed, or 1–3-pinnatisect, or palmately lobed, sometimes trifoliolate, pinnately,
palmately, or bipinnately compound; cauline leaves alternate, rarely opposite or whorld, simple
or compound as basal leaves, petiolate or sessile and base cuneate to attenuate or auriculate to
sagittate, margin entire, dentate, or variously lobed. Racemes several to many-flowered,
ebracteate or rarely bracteate throughout, corymbose or in panicles, elongated in fruit; rachis
straight or rarely flexuous; fruiting pedicels slender or thickened, erect, divaricate, or reflexed.
Sepals ovate or oblong, free, caducous or rarely persistent, erect to rarely spreading, equal or
unequal, base of lateral pair saccate or not. Petals white, pink, purple, or violet, never yellow,
erect to spreading, longer than sepals, rarely absent; blade obovate, spatulate, oblong, or
oblanceolate, apex obtuse or emarginate; claw absent or strongly differentiated from blade,
longer to shorter than sepals, glabrous, unappendaged, entire. Stamens 6 and slightly to strongly
tetradynamous, rarely 4 and equal in length, erect or spreading, exserted or included; filaments
wingless, unappendaged, glabrous, free, dilated or not at base; anthers ovate to oblong or linear,
not apiculate at apex. Nectar glands confluent and subtending bases of all stamens; median
nectaries 2 or rarely 4 or absent; lateral nectaries annular or semiannular. Ovules 4−80 per ovary;
placentation parietal. Fruit dehiscent capsular siliques, linear or rarely oblong, elliptic, or
narrowly lanceolate, slightly to strongly latiseptate, not inflated, unsegmented; valves papery, not
veined or rarely proximal part with an obscure midvein, glabrous or very rarely hairy, not keeled,
flat, not covering entire fruit width, smooth or torulose, wingless, unappendaged, dehiscing
elastically acropetally, spirally or circinately coiled; gynophore absent; replum strongly
flattened, visible; septum complete, membranous, tansluscent, veinless; style distinct and
slender, rarely obsolete, persistent, glabrous; stigma capitate, entire, unappendaged. Seeds
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uniseriate, wingless or not margined, oblong to ovate, slightly flattened; seed coat smooth,
minutely reticulate, colliculate, or rugose; mucilaginous or not when wetted; cotyledons
accumbent or very rarely incumbent. × = 7, 8.
256 species: worldwide and with native species on all continents except Antarctica.
References:
Interactive key to the genera is being prepared
2
acris
africana
altigena
anemonoides
angulata
angustata
anhuiensis
amara
apennina
appendiculata
arakiana
armoracioides
asarifolia
astoniae
auriculata
balnearia
bellidifolia
bilobata
blaisdellii
bodinieri
bonariensis
breweri
bulbifera
bulbosa
calcicola
caldeirarum
californica
calthifolia
carnosa
caroides
castellana
changbaiana
chelidonia
chenopodiifolia
cheotaiyienii
chilensis
circaeoides
clematitis
concatenata
conferta
constancei
cordata
cordifolia
corymbosa
crassifolia
debilis
delavayi
densiflora
dentata
depressa
digitata
diphylla
dissecta
douglasii
elegantula
engleriana
enneaphyllos
enriquei
eremita
fallax
fargesiana
flagellifera
flexuosa
fragariifolia
franchetiana
franklinensis
fulcrata
gallaecica
georgiana
geraniifolia
glacialis
glanduligera
glauca
glechomifolia
gouldii
gracilis
graeca
granulifera
granulosa
griffithii
gunnii
heptaphylla
hirsuta
hispidula
holmgrenii
hydrocotyloides
hygrophila
iliciana
impatiens
jamesonii
jejuna
jonselliana
keysseri
kitaibelii
komarovii
kruesselii
lacustris
latior
lazica
leucantha
lihengiana
lilacina
lineariloba
lojanensis
longii
longipedicellata
loxostemonoides
luxurians
lyrata
macrocarpa
macrophylla
majovskii
marginata
maritima
matthioli
maxima
mexicana
micranthera
microphylla
microthrix
microzyga
moirensis
monteluccii
multiflora
multijuga
nepalensis
niigatensis
nipponica
nuttallii
nymanii
obliqua
occidentalis
oligosperma
opizii
ovata
pachystigma
papillata
papuana
parviflora
pattersonii
paucifolia
paucijuga
pectinata
pedata
penduliflora
pensylvanica
pentaphyllos
penzesii
picta
plumieri
pratensis
prorepens
pulchella
Species in red are not yet described
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purpurascens
purpurea
quinquefolia
raphanifolia
repens
resedifolia
rivularis
robusta
rockii
rostrata
rotundifolia
rupicola
scaposa
schinziana
schulzii
scutata
seidlitziana
seravschanica
silana
simplex
speciosa
sphenophylla
stellata
stenoloba
subcarnosa
tanakae
tangutorum
tenera
tenuifolia
tenuirostris
tianqingiae
trichocarpa
trifida
trifolia
trifoliolata
tryssa
tuberosa
uliginosa
uniflora
umbellata
variabilis
victoris
violacea
volckmannii
vulgaris
waldsteinii
yezoensis
yunnanensis
Cardamine acris Grisebach, Spicil. Fl. Rumel. 1: 253. 1843; C. raphanifolia Pourret subsp.
acris (Grisebach) O. E. Schulz, Bot. Jahrb. Syst. 32: 512. 1903. TYPE: [F.Y.R. Makedonija]
“gregarie ad fontem m. Kobelitza [Kobelica] juxta Mandram Weitzensem alt. 4200′, 21 Jul
1839, Grisebach 943 (lectotype designated by Strid (1986: 257), GOET; isolectotype, K).
Cardamine africana Linnaeus, Sp. Pl. 2: 655. 1973. Lectotype (designated by Marais, 1970):
Hermann, Paradisus Batavus Fig. p. 202 (1698).
Cardamine innovans O. E. Schulz, Bot. Jahrb. Syst. 32: 417. 1903. TYPE: Guatemala, above
Tecpam, 2500 m, 1882, F. C. Lehmann 1475 (lectotype, here designated, G-BOIS).
Cardamine borbonica Persoon, Syn. Pl. 2: 195. 1807. TYPE: Réunion [Bourbon], Bory de St.
Vincent (holotype, G).
Cardamine holtziana Engler & O. E. Schulz, Bot. Jahrb. Syst. 32: 416. 1903. TYPE: Tanzania,
Lushoto District, W. Usambara Mts, between Wambugu and Malalo, A. Engler 1356
(holotype, B).
Cardamine ovata Bentham var. corymbosa Britton, Bull. Torrey Bot. Club 16: 16. 1889. TYPE.
Bolivia, [La Paz], Undavi, 10000 ft, Oct 1885, H. H. Rusby 1206 (holotype, NY!).
Herbs, perennial. Rhizomes slender, not tuberous. Stems (5−)25−70(−80) cm, erect to
ascending, simple or sometimes branched near middle, glabrous throughout or pubescent. Basal
leaves not rosulate; lower and middle cauline leaves 4−10(−15) cm, trifoliolate; petiole 2−6(−7)
cm; terminal leaflet (1−)1.5−7 × 1.5−4 cm, lanceolate to ovate, sparsely strigose with trichomes
to 0.6 mm, base cuneate to subobtuse, margin ciliolate or glabrous, serrate or crenate, teeth
mucronate, apex acute, petiolule 2−2.5 cm; lateral leaflets same size or slightly smaller than
terminal one, base oblique. Racemes ebracteate, slightly elongated or not elongated in fruit;
rachis straight; fruiting pedicels 0.5−3 cm, ascending, stout, straight. Sepals oblong, 1.5−4 ×
1−1.5 mm, not saccate, caducous, glabrous; petals white or lavender, spatulate, 3.5−9 × 2−4 mm,
apex obtuse; filaments 2.5−5 mm; anthers oblong, 1−1.5 mm; ovules 8−20 per ovary. Fruits
linear, (2.5−)3.5−5(−6) cm × 1.5−2.5 mm, glabrous; style 0.5−1.5 mm, stout. Seeds brown,
oblong, 1.5−2.5 × 1−1.5 mm.
Flowering: Sep−Jan.
Habitat: Moist or wet areas, mixed forests, disturbed places, roadside clearings, damp shady
areas.
Elevation: 500−3400 m.
Distribution: native of Africa (Comores, Ethiopia, Kenya, Madagascar, South Africa, Tanzania,
Ugunda), naturalized in Asia (India, Indonesia, Malesia, Papua New Guinea, Sri Lanka), Central
America (Costa Rica, El Salvador, Guatemala, Honduras, Mexico [Chiapas]), and South
America [Argentina (Misiones, Salta, Tucumán), Bolivia (Cochabamba, La Paz, Santa Cruz,
Tarija), Brazil (Parana, Río Grande Do Sul, Santa Catalina), Columbia (Cundinamarca),
Ecuador (Santiago-Zamora), Peru (Amazonas, Cajamarca, Cuzco), Venezuela (Mérida)].
Specimens examined: ARGENTINA. Misiones: Gral. Manuael Belgrano, 8 km de Bernardo de
Irigoyen hacia San Antonio, 26°12′S, 53°40′W, Morrone et al. 1378 (MO, SI). Salta: Santa
Victoria, camino al Angosto del Baritú, 22°30′S, 64°45′W, Ahumada & Agüero 8343 (SI); Parq
Nac. Baritú, Lipeo, Rio Naranjo, Zuloaga et al. 1165 (BACP, SI); Anta, Parque Nac. El Rey,
Picada al Chomo Los Loios, Ezcurra, Ponce & Brown 431 (SI). Dep. Anta, Res. Nac. Fca. Del
Rey, Ezcurra et al. 431 (BACP). Tucumán: Dep. Chicligasta, arroyo Celeste, R 65, Km 28,
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27°20′S, 65°50′W, Boelcke et al. 5474 (BAA). BOLIVIA. Chuquisaca. Prov. Hernando Siles,
Comunidad Vallecito, Villalobos & Jimenez 903 (MO). Cochabamba: Chapare, Cochabamba 95.2 km
hacia Villa Tunari, Beck 7285 (BACP, GH, MO). La Paz: Yungas, Bang 227 (BM, E, K, MO,
NY, W); Murillo, 44 km below Lago Zongo, 16°03′S, 68°01′W, Solomon 10777 (GH, MO),
Cahua, Valle de Zongo, Beck & Foster 13949 (LPB, MO); Nor Yungas, 17.3 km NE Chuspipata
on road to Yolosa, 16°13′S, 67°47′W, Solomon 12547 (GH, MO, NY); 4.5 km below Yolosa,
then 10 km W on rd up the Rio Huarinilla, 16°12′S, 67°50′W, Solomon 8551 (GH, MO). Sant
Cruz: Caballero, Enpalme, 17°49′S, 64°36′W, Rivera et al. 63 (MO). Tarija: Arce, Rio
Chillaguatas, on trail between Sidaras and Tariquia, 22°05′S, 64°25′W, Solomon 11270 (GH,
MO); entre Emboruzú y La Mamora, Zuloaga et al. 1058 (BACP, SI). BRAZIL. Parana: Tres
Barras, Dusén 17565 (GH, NY, S). Rio Grande Do Sul: Morro do Forno-Josafá, Sobral &
Jarenkow 8022 (F). Santa Catalina: Sun. Sao Miguel d’ Oeste, forest above Rio Reperi-guacu,
Peperi, 26°32′S, 53°44′W, Smith & Reitz 12783 (B, F, GH, MO, NY, P); Bom Retiro, Agua Boa,
Riozinho, Smith & Kline 7892 (GH). COLOMBIA. Cundinamarca: Quetame to Susumuco,
Pennell 1344 (NY) ECUADOR. Andes, without locality, Spruce 5378 (E, G, GH, NY, P, W).
Santiago-Zamora: Río Itzintza, 2°40′S, 78°W, Camp E-1218 (GH, NY); above Tambo
Valladolid, Steyermark 54625 (NY). PERU. Amazonas: Pongará, Yambrasbamba, Tillett 673364 (F, GH). Cajamarca: San Ignacio, San José de Lourdes, 4°59′S, 78°54′W, Díaz et al. 10540
(MO). Cusco: La Convención, Santa Ana, Madre Selva, 12°53′S, 72°45′W, Valenzuela et al.
3047 (MO); Valle de Santa Ana, above Quillabamba, Plowman & Davis 4803 (GH).
VENEZUELA. Merida: La Mucuy, Bernardi 1835 (NY).
Both Schulz (1903) and Rollins (1993) recognized Cardamine africana and C. innovans as
inDep. endent species, but the alleged differences in the presence vs. absence of leaf marginal
trichomes are unreliable.
Cardamine altigena O. E. Schulz, Bot. Jahrb. Syst. 62: 479. 1929. TYPE:
Herbs, perennial, glabrous or sparsely pubescent. Rhizome slender. Stems 2–5 dm, ascending
to prostrate, rooting from lower nodes. Leaves 3–7-foliolate or rarely unifoliolate, glabrous or
ciliate; basal soon withered, 2–7 cm; cauline leaves remote, few, to 6 cm, the petiole not
auriculate and to 2 cm; terminal leaflet trifid, orbicular to reniform becoming oblanceolate in
uppermost leaves, 0.5–1.5(–3) × 0.1–1.2(–2) cm, sessile or with a petiolule 1–8 mm; lateral
leaflets trifid to oblanceolate, 2–7 × 1–5 mm, petiolule ca. 1 mm. Racemes ebracteate, 3–10flowered; fruiting pedicels erect to divaricate, straight, 8–15 mm. Sepals oblong, 1.8–2.7 mm;
petals white, spatulate, 3.5–6 × 1–2 mm, apex obtuse; stamens 6, filaments subulate, anthers 0.3–
0.5 mm; ovules – per ovary. Fruits linear, 1.3–3.5 cm × 1.3–1.8 mm; style 0.4–2 mm. Seeds
oblong, 1.4–1.7 × ca. 1 mm, smooth.
Habitat: wet grassy slopes, near running water, along streams.
Elevation: 1700–4000 m.
Distribution: New Guinea.
Cardamine anemonoides O. E. Schulz, Bot. Jahrb. Syst. 32: 340. 1903; Dentaria corymbosa
Matsumura, Bot. Mag. (Tokyo) 13: 52. 1899; Cardamine matsumurana Nemoto, Fl. Japan
Suppl. 263. 1936; not C. corymbosa J. D. Hooker, Icon. Pl. 7: 686. 1844. TYPE: Japan, select
one of the five syntypes. (lectotype, here designated, TI!).
Cardamine anemonoides var. suavis O. E. Schulz, Bot. Jahrb. Syst. 32: 341. 1903. TYPE:
Schikoku, Prof. Tosa, pr. Nanokawa, 17 Apr 1890, Watanabe s.n. (holotype, GH!).
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Herbs, perennial. Rhizomes creeping, scaly, subfleshy, without tuberous knots, 1–3 mm in
diam., not stoloniferous. Stems, 6–30 cm, slender, erect, simple, glabrous. Rhizomal leaves
withered by flowering time; cauline leaves 2–5, trifoliolate or very rarely simple, not auriculate,
hirsutulous with antrorse trichomes 0.3–0.5 mm; petiole (0.5–)1–3.5 cm; terminal leaflet (or leaf
when simple) broadly ovate, lanceolate, or elliptic, rarely obovate, 1–6 × 0.6–2.5 cm, on a
petiolule to 1 cm, base cuneate, margin ciliolate antrorse trichomes to 0.1 mm, crenate-serrate to
serrate, the teeth mucronate, apex acute to acuminate; lateral leaflets 1 pair, sessile, similar to
terminal leaflet in shape but smaller and often with oblique base. Racemes ebracteate, 2–7flowered; fruiting pedicels ascending to suberect, 1–2.5 cm, straight, slender. Sepals narrowly
oblong, 3–4 × 1–1.5 mm; petals white, narrowly obovate, 6–10 × 2–3 mm, apex rounded; median
filament pairs 5–6 mm, lateral pair 4–5 mm; anthers oblong, 1.2–1.5 mm; ovules 8–20 per ovary.
Fruit linear, 3–4.7 cm × 1–1.5 mm; valves smooth, glabrous; style 3–8 mm. Seeds brown,
oblong, 1–15 × 0.7–1 mm.
Flowering: Apr–May.
Habitat: mountain forests.
Elevation: 500–1300 m.
Distribution: Japan (Honshu).
Cardamine angulata W. J. Hooker, Fl. Bor.-Amer. 1: 44. 1829. TYPE :
Cardamine angulata var. alba Nuttall, in Torrey & A. Gray, Fl. N. Amer. 1: 84. 1838. TYPE:
Cardamine angulata var. hirsuta O. E. Schulz, Bot. Jahrb. Syst. 32: 407. 1903. TYPE:
Cardamine angulata var. pentaphylla O. E. Schulz, Bot. Jahrb. Syst. 32: 407. 1903. TYPE:
Cardamine helleriana O. E. Schulz, Bot. Jahrb. Syst. 32: 547. 1903. TYPE:
Dentaria grandiflora Rafinesque, Atlantic J.-Extra of No. 6 (Herb. Raf.) 47. 1833. TYPE:
Herbs, perennial, glabrous or sparsely pubescent. Rhizomes slender, cylindrical, to 2 mm in
diam.; stolons absent. Stems (1.5–)2.5–8.5(–10) dm, erect, simple, sparsely to densely hirsute at
base. Rhizomal leaves 3(or 5)-foliolate, (4–)7–20(–22) cm; petiole (2–)4–12(–14) cm; terminal
leaflet ovate to broadly lanceolate, 1.5–7(–9) cm, with a petiolule (0.3–)0.5–1.5 cm, base
cuneate, rarely subreniform or obtuse, margin 3–5(–7)-lobed or -toothed, puberulent; lateral
leaflets about as large as or smaller than terminal leaflet, subsessile, margin same as terminal
leaflet; cauline leaves (3 or)4–8, 3(or 5)-foliotate; petiole 1–4 cm, base not auriculate; terminal
lobe broadly ovate to narrowly lanceolate, 2–7 × 0.6–4 cm, minutely pubescent along margin,
petiolulate or sessile; lateral leaflets similar to terminal one but smaller, sessile, margin dentate
or rarely entire. Racemes ebracteate; fruiting pedicels ascending to divaricate, (0.9–)1.2–2.5 cm.
Sepals oblong, 2.5–4 × 1.3–2 mm, erect, base of lateral pair saccate; petals white or rarely
pinkish, obovate, 8–15 × 4–8 mm, clawed, apex rounded or emarginate; median filament pairs
3.5–6 mm, lateral pair 2–3.5 mm; anthers oblong, 0.8–1.2 mm. Fruit linear, 1.5–3.2 cm × 1.4–2
mm; valves glabrous; style (0.5–)1–4 mm; ovules and seeds 10–16 per fruit. Seeds dark brown,
oblong, 1.8–2.3 × 1–1.2 mm. 2n = 40.
Flowering; Apr–Jun.
Habitat: moist grounds, streambanks, swampy or damp woods, thickets, wet meadows.
Elevation: 0–900 m.
Distribution: Canada (British Columbia), United States (Alaska, N California, Oregon,
Washington).
2. Cardamine angustata O. E. Schulz, Bot. Jahrb. Syst. 32: 349. 1903; Dentaria heterophylla
Nuttall, Gen. N. Amer. Pl. 2: 66. 1818; Cardamine heterophylla (Nuttall) A. Wood Amer.
Bot. & Fl. 38. 1870; not Host, Syn. P. Austral. 366. 1797; not Lapeyrouse, Hist. Abr. Pl. Pyr.
6
377. 1813; not Bory, Ann. Sci. Gen. Phys. 3: 6. 1820; not W. J. Hooker, Comp. Bot. Mag. 1:
273. 1835. TYPE:
Cardamine angustata var. ouachitana E. B. Smith, Brittonia 34: 379. 1982. TYPE: United
States, Arkansas, Polk Co., common alon the Cossatot River, 12.1 miles S of the junction of
Hwys 375 and 8 near Mena, 19 Mar 1982, E. B. Smith 3664 (holotype, UARK; isotypes M)!,
NY!, US!).
Herbs, perennial, glabrous or rarely sparsely pubescent. Rhizomes fleshy, moniliform,
segments fusiform, 3–6 mm in diam.; stolons absent. Stems 1.2–3(–4) dm, erect, simple,
glabrous or pubescent. Rhizomal leaves 3-foliolate, to 24 cm; petiole (3–)5–12(–16) cm;
terminal leaflet broadly ovate to rhombic-obovate, 1.5–6(–8) cm, with a petiolule (0.2–)0.5–1.5(–
2) cm, base cuneate, rarely subtruncate, margin coarsely dentate to crenate or 3-lobed, puberulent
or not; lateral leaflets about as large as or smaller than terminal leaflet, subsessile or with a
petiolule 2–10 mm, margin same as terminal leaflet; cauline leaves 2(or 3), 3-foliolate, alternate
or rarely opposite, different in morphology from rhizomal leaves; petiole 0.5–2 cm, base not
auriculate; terminal leaflet narrowly lanceolate to narrowly oblong or oblong-lanceolate, 2–7 cm
× 3–6 mm, minutely puberulent along margin, petiolulate or sessile; lateral leaflets similar to
terminal one but smaller, sessile, margin dentate or rarely entire. Racemes ebracteate; fruiting
pedicels ascending to divaricate, 1.5–4 cm. Sepals oblong, 5–7.5 × 1–2 mm, erect, base of lateral
pair slightly saccate; petals purple to pale pink, oblanceolate, 9–18 × 2–5 mm, clawed, apex
rounded; median filament pairs 5–10 mm, lateral pair 3.5–8 mm; anthers linear, 1.5–3 mm. Fruit
linear, 2.5–4 cm × 1.5–2.5 mm; valves glabrous; style (5–)7–11 mm; ovules and seeds 8–12 per
fruit. Seeds dark brown, oblong, 2–2.5 × 1–1.5 mm. 2n = ca. 128.
Flowering: Mar–May.
Habitat: moist woods, wooded ridges and bottomlands, flood plains, shady ravines, stream beds.
Elevation: 300–1300 m.
Distribution: United States (Alabama, Arkansas, Delaware, District of Columbia, Georgia,
Indiana, Kentucky, Maryland, Mississippi, New Jersey, North Carolina, Ohio, Pennsylvania,
South Carolina, Tennessee, Virginia, West Virginia).
Cardamine anhuiensis D. C. Zhang & J. Z. Shao, Bull. Bot. Res., Harbin 6(2): 127. 1986.
TYPE. China, Anhui, Tongling Xian, Lang-Keng, 200 m, 14 Apr 1985, Shao Jianzhang
84050 (holotype, ANUB; isotypes, NAS!, PE!).
Cardamine jinshaensis Q. H. Chen & T. L. Xu, Acta Bot. Yunnan. 15: 361. 1993. TYPE: China,
Guizhou, Jincha Ciann, Qingmen, 1000 m, 26 Apr 1988, Qianxi Expedition 330 (holotype,
HGAS).
Herbs, perennial, sparsely pilose to glabrous. Rhizomes stout, often without stolons. Stems
11–35 cm, erect, simple or sometimes 1- or 2-branched above, not flexuous. Basal leaves
trifoliolate or simple; petiole 2–17 cm; terminal leaflet or leaf blade suborbicular or rarely ovateorbicular or reniform-orbicular, 0.6–3.5 × 1–5 cm, base cordate or rarely subtruncate, margin
crenate or rarely repand-crenate; petiolule of terminal leaflet 0.7–1.4 cm, that of lateral leaflets
much shorter; lateral leaflets similar to terminal but much smaller; cauline leaves 3(or 5)foliolate; petiole 1.3–3.5 cm, base not auriculate; terminal leaflet ovate-orbicular or subreniform,
1.5–3 × 1–3.2 cm, with a petiolule to 6 mm, margin crenate or crenate-repand; lateral leaflets
similar to terminal one, shortly petioulate to subsessile. Racemes ebracteate; fruiting pedicels
divaricate to divaricate-ascending, 0.6–1.5 cm, slender. Sepals oblong, 2–2.8 × 0.8–1.2 mm, base
not saccate; petals white, spatulate, 4–6 × 1.5–2.5 mm, apex rounded; median filament pairs 2.5–
4 mm, lateral pair 1.5–2.5 mm; anthers oblong, 0.5–0.7 mm; ovules 24–30 per ovary. Fruit
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linear, 2–4 cm × 1–1.5 mm; valves smooth, glabrous; style 1–3 mm. Seeds brown, oblong, 1.2–2
× 0.8–1.3 mm, wingless.
Flowering: Mar–May.
Habitat: shady slopes, along ditches.
Elevation: 0–1000 m.
Distribution: China (Anhui, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Zhejiang).
Cardamine amara Linnaeus, Sp. Pl. 2: 656. 1753. TYPE: Herb. Linn. No. 835.17 (lectotype
designated by Khatri (1989: 92), LINN).
Cardamine apennina Lihová & Marhold, Pl. Syst. Evol. 245: 85. 2004. TYPE:
Cardamine appendiculata Franchet & Savatier, Enum. Pl. Japon. 2: 281. 1879; C. macrophylla
Willdenow var. appendiculata (Franchet & Savatier) Yatabe, Bot. Mag. (Tokyo) 6: 100. 1892;
Dentaria appendiculata (Franchet & Savatier) Matusmura, Bot. Mag. (Tokyo) 31: 51. 1899.
TYPE: Northern Japan, Savatier s.n. (holotype, P!).
Herbs, perennial, Rhizomes creeping, not scaly, slender, without tuberous knots, 1–4 mm in
diam., not stoloniferous. Stems (20–)30–65 cm, stout or slender, erect, simple, glabrous, slightly
flexuous. Rhizomal leaves 1–5 cm; petiole 2–7 cm; terminal leaflet oblong to ovate, (1–)2–4 ×
0.8–2 cm, sessile or petiolulate, base cuneate, margin serrate to crenate, apex acute; lateral
leaflets 1 or 2 pairs, similar to terminal but smaller; cauline leaves (3–)5–10, glabrous or
scabrous with antrorse trichomes to 0.2 mm; petiole (1–)2–7(–10) cm, basal auricles 1–6 × 0.5–2
mm; terminal leaflet broadly ovate, lanceolate, elliptic, or oblong, (1.5–)3–7(–10) × (0.5–)1–4(–
7) cm, glabrous or pubescent, subsessile or petiolulate, base cuneate, margin ciliolate with
appressed trichomes to 0.2 mm, crenate, serrate, or serrulate, the teeth mucronate, apex acute,
rarely subacuminate; lateral leaflets 2–5 pairs, sessile, similar to terminal leaflet. Racemes
ebracteate, 10–30-flowered; fruiting pedicels divaricate, 1–2(–2.7) cm, straight, slender. Sepals
oblong, 2.5–3.5 × 1–1.5 mm; petals white, narrowly obovate, 7–9 × 3–4 mm, apex rounded;
median filament pairs 4–5 mm, lateral pair 3–4 mm; anthers oblong, 1.2–1.5 mm; ovules 8–16
per ovary. Fruit linear, 2–3 cm × ca. 1.5 mm; valves smooth, glabrous; style (2–)3–6 mm. Seeds
brown, ovoid, 1.5–2.5 × 1–1.5 mm.
Flowering: May–Jul.
Habitat: wet areas along streams, shady places in forests.
Elevation: 200–2000 m.
Distribution: Japan (Hondo, Honshu).
Cardamine arakiana Koidz, Acta Phytotax. Geobot. 5: 119 (1936). TYPE:
Herbs, perennial. Rhizomes slightly thickened. Stems (10–)15–35(–42) cm; erect pubescent
or rarely glabrous, usually branched above. Basal leaves simple, trifoliolate or rarely 5-foliolate,
6–17 cm; blade or terminal leaflet orbicular- or cordate-reniform, 1.3–4.5(–5.5) cm, 1.5–5.5(–
6.5) mm wide, base cordate, margin repand or crenate, apex rounded; lateral leaflets
suborbicular, 0.5–1 cm, 0.5–1.3 mm wide, base truncate to cordate, margin crenate; cauline
leaves trifoliolate or rarely 5-foliolate, 2.5–15 cm, adaxially glabrous, pubescent adaxially;
terminal leaflet ovate to suborbicular, 1–2.5 cm, 1–3 cm wide, base cordate to obtuse, margin
crenate; lateral leaflets elliptic, ovate, or orbicular, 0.5–2.5 cm, 0.3–2 cm wide, usually
petiolulate. Racemes ebracteate, 2–10 cm in fruit; fruiting pedicels 1–2 cm, ascending. Sepals 2–
8
2.5 mm, ca. 1 mm wide, sparsely pubescent; petals white, 5–6 mm, 2–3 mm wide; filaments 2–3
mm. Fruits linear, 2.5–4 cm, ca. 1 mm wide, glabrous; style 2–3 mm. Seeds 1.5–2 mm. 2n = 32.
Flowering: Mar–Apr.
Habitat: Grassy silty-clay forest floor, forest margins in low mountains.
Distribution: Japan (Honshu).
Cardamine armoracioides Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 27(2, 4): 293.
1854 (1855). TYPE: Venezuela, New Grenada, Linden 1416 (holotype, KW?; isotypes, BM!,
G!, K!, P!).
Cardamine lanceolaris Linden & Plancheon, Trois. Voy. Linden, Bot., Pl. Columb. 1: 12. 1863.
TYPE: Same as that of C. armoracioides but holotype at P!.
Herbs, perennial, with woody base, sometimes annual. Rhizomes absent. Stems 20−50 cm,
erect or decumbent, simple or branched above, glabrous. Cauline leaves (lower and middle ones)
4−22 cm, simple or rarely with a tiny lateral leaflet; petiole (1.5−)3−7 cm; leaf blade (3−)4.5−15
× (1.5−)2−5.5 cm, lanceolate to ovate, glabrous or sparsely pubescent with trichomes to 0.5 mm,
base cuneate, margin ciliolate with trichomes to 0.2 mm, serrate, teeth mucronate, apex long
acuminate or rarely acute; uppermost leaves smaller, reduced into bracts. Racemes bracteate at
least along proximal half; bracts simple, linear to linear-oblanceolate, petiolate, reduced in size
upward; fruiting pedicels 1−2 cm, divaricate to ascending, stout, straight. Sepals oblong, 2.5−3.5
× 1−1.5 mm, not saccate; petals white, spatulate to oblanceolate, 4.5−6 × 1.5−2.5 mm; filaments
3−4 mm; anthers oblong, 0.7−0.9 mm; ovules 10−14 per ovary. Fruits 3−5.5 cm × 1.5−2 mm;
style 1−3 mm. Seeds brown, oblong, 2−2.7 × 0.9−1 mm.
Flowering: May−Sept.
Habitats: Rich steep forest slopes.
Elevation: 2100−2700 m.
Distribution: Colombia?, Venezuela (Mérida).
Specimens examined: COLOMBIA. Jaji, Moritz 1052 (B). VENEZUELA. New Grenada,
Linden 1414 (BM). Merida: above La Isla, above Tabay, Steyermark 56596 (F, GH); La Isla,
arriba de La Mucuy, 8 km arribay de Tabay, Bunting 4540 (F, GH); Laguna de Coromoto,
June−Sept. 1958, Schwabe s.n. (B); Mucuy above Merida, 31 Dec. 1972, Schuabe s.n. (B).
The locality from Colombia needs further checking to see if it is indeed in that country or in
Venezuela.
Cardamine asarifolia Linnaeus, Sp. Pl. 2: 654. 1753. TYPE: Hermann, P. Paradisus batavus,
1698, p. 203, entitled “NASTURTIUM montanus Asari folio” [lectotype designated by
Marhold (1996: 114)].
Cardamine astoniae I. Thompson, Muelleria 9: 156. 1996. TYPE: Australia, Victoria,
Snowfields, Bogong High Plains, between Rocky Valley Reservoir and Basalt Hill, Falls
Creek area, 36º54′S, 147º18′E, 1650 m, 28 Dec 1994, Ian Thompson 84 (holotype, MEL).
Herbs, perennial, glabrous throughout. Rhizomes slender, sometimes branched. Stems to 25
cm, erect, simple. Basal leaves not rosulate, sometimes clustered at stem base, somewhat fleshy,
long petiolate, to 15 cm, simple or pinnately compound and 3−5-foliolate; terminal leaflet ovate
to elliptic, cuneate to cordate at base; lateral leaflets smaller, orbicular to elliptic; cauline leaves
several, petiolate, not auriculate at base, pinnately compound or pinnatisect, lateral
segments/leaflets strongly angled forward. Racemes ebracteate, many flowered, short; fruing
9
pedicels 10−20 mm. Sepals ovate, 3−4 mm; petals white or pink outside, differentiated into
blade and claw, 6−11 × 3−6 mm; stamens 6. Fruits linear, erect to suberect, 2−3 cm × 2−2.5
mm. Seeds oblong-elliptic, ca. 2 mm.
Flowering: Nov−Jan.
Habitat: among tussocks, moist to boggy alpine to subalpine areas.
Elevation: ca. 1650 m.
Distribution: Australia (New South Wales, Tasmania, Victoria).
Cardamine auriculata S. Watson, Proc. Amer. Acad. Arts 17 : 319. 1882. TYPE:
Herbs, annual, sparingly hispid near base. Stems 1.5−4 dm, erect to ascending or decumbent,
slender, branched. Leaves pinnately compound, with 2 or 3 pairs of leaflets, petiolate, not
auriculate; leaflets petiolulate, ovate to ovate-oblong, 1−2 cm, crenate-dentate, sparsely
pubescent or glabrous, minutely ciliate or not, sometimes a subsidiary leaflet present at lower
side of rachis. Racemes ebracteate; fruiting pedicels divaricate-ascending, 6−10 mm, glabrous.
Sepals 2.5−3.5 mm, not saccate; petals white, spatulate, 6−8 mm; ovules −. Fruits linear, 2.5−4
cm × 1.5−2 mm; style 1.5−3 mm, slender. Seeds oblong, ca. 1.5 mm × 1 mm, occasionally
winged.
Flowering: Apr−Jul.
Habitat: limestone ledges, wet banks, near streams, deep canyons, spring, wooded slopes.
Distribution: Mexico (Hidalgo, Nuevo León, San Luis Potosí, Tamaulipas).
Cardamine balnearia Standley & Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23: 157. 1944.
TYPE: Guatemala, Volcán de Zuni, ca. 2450 m, P. Standley 83332 (holotype, F!; isotype, US!).
Herbs, perennial, glabrous throughout. Rhizomes thickened, erect. Stems 3−3.5 dm, erect,
simple below, branched and densely leafy above base. Leaves ca. 15 cm, 9−13-foliolate; petiole
slender, 4−7 cm, not auriculate at base; lateral leaflets 1−3 cm × 4−12 mm, broadly ovate to
oblong or oblong-lanceolate, glabrous, on a petiolule 4−6 mm, base oblique and cuneate to
rounded, margin dentate to slightly lobed, the teeth mucronate, apex acute; terminal leaflet
similar to lateral ones but slightly larger. Racemes ebracteate or only lowermost flowers
bracteate; fruiting pedicels 1−1.5 cm, ascending, straight, slender. Sepals 2.5−3 × ca. 1 mm,
purplish, oblong; petals 5−6 × ca. 2 mm, white tinged with purple; filaments 4−5 mm; anthers ca.
1 mm; ovules −. Fruits (immature) ca. 3.5 cm × 1 mm; style ca. 3 mm. Seeds not seen.
Flowering: Jan.
Habitat: wet mossy banks.
Elevation: ca. 2450 m.
Distribution: Guatemala.
Notes: known only from the type collection.
Cardamine bellidifolia Linnaeus, Sp. Pl. 2: 654. 1753. TYPE: Herb. Linn. 835.1. (lectotype
designated by Khatri (1990a: 442), LINN).
Cardamine bellidifolia var. beringense A. Porsild, Proc. Trans. Roy Soc. Canada 32: 31. 1938.
TYPE:
Cardamine bellidifolia var. laxa Lange, Meddel. Gronland 3: 251. 1887. TYPE:
Cardamine bellidifolia var. pachyphylla Coville & Leiberg, Proc. Biol. Soc. Wash. 11: 170.
1897. TYPE:
10
Cardamine bellidifolia var. pinnatifida Hultén, Madroño 19: 223. 1968. TYPE:
Herbs, perennial, glabrous throughout, somewhat cespitose. Rhizomes and stolons absent;
caudex with slender or rarely stout, rhizome-like branches. Stems 0.1–0.8(–1.4) dm, erect to
ascending, many, glabrous, subscapose. Basal leaves simple, rosulate, (0.6–)1.2–5(–7) cm;
petiole (3–)1–3.5(–5.5) cm; leaf blade ovate to oblong, rarely oblanceolate to obovate, (0.4–)0.8–
1.7(–2.5) × (0.2–)5–1(–1.6) cm, base cuneate to obtuse, margin entire or rarely repand or
obtusely small toothed, not puberulent, apex obtuse; cauline leaves absent or 1(or 2), simple,
short petiolate, base not auriculate, similar in shape and much smaller than basal leaves.
Racemes ebracteate; fruiting pedicels ascending to erect, 3–6(–8) mm. Sepals oblong, 2–3(–4) ×
0.8–1.5(–2) mm, erect, base of lateral pair not saccate; petals white, oblanceolate, 4–5.5(–7) ×
1.3–2(–2.5) mm, not clawed, apex rounded or emarginate; median filament pairs 2.5–4 mm,
lateral pair 2–3 mm; anthers oblong, 0.5–0.9 mm. Fruit linear, (0.8–)1.3–2.8(–3.7) cm × 1.3–2
mm; valves glabrous; style 0.5–3 mm; ovules and seeds 8–18 per fruit. Seeds brown, oblong,
1.5–2 × 0.9–1.2 mm. 2n = 16.
Flowering: Jun–Sep.
Habitat: mossy places, tundra, marshes at stream headwaters, cliffs, talus slopes, barren chert
slopes, mosit rock crevices, rocky slopes, alpine brooks, sandy beaches, moist rocky stream beds.
Elevation: 0–2300 m.
Distribution: Canada (Alberta, British Columbia, Labrador, Nunavut, Northwest Territories,
Quebec, Yukon), Greenland, n Europe, Russia (Siberia, Far East), United States (Alaska,
California, Maine, New Hampshire, Oregon, Washington).
Cardamine bilobata Kirk, Stud. Fl. 27. 1899. TYPE: New Zealand, Kurow, Petrie s.n.
(holotype, W).
Cardamine blaisdellii Eastwood, Bot. Gaz. 33: 146. 1902; C. microphylla Adams subsp.
blaisdellii (Eastwood) D. F. Murray & S. Kelso; C. microphylla var. blaisdellii (Eastwood)
Khatri. TYPE:
Herbs, perennial, glabrous throughout or rarely pilose. Rhizomes slender, cylindrical, 0.7–1.5
mm in diam.; stolons absent. Stems 0.5–2(–2.5) dm, erect or ascending, simple, glabrous or
rarely pilose. Rhizomal leaves pinnately 5–7-foliolate, not fleshy, 2.5–12 cm; petiole 1.5–4(–9)
cm; terminal leaflet suborbicular to broadly obovate, 4–15 × 2.5–14 mm, glabrous or rarely
pilose, base obtuse to subcordate, margin 3–5-toothed, apex apiculate, petiolule 1.5–3 mm;
lateral leaflets toothed or rarely entire, about as large as or smaller than terminal leaflet, similar in
other aspects; cauline leaves 1–3, (3–)5-foliotate, alternate; petiole 0.2–2(–6.5) cm, base not
auriculate; terminal leaflet obovate to oblanceolate, 5–15 × 3–10 mm, subsessile or with a
petiolule to 2 mm, base cuneate, margin 3-toothed or entire, apex apicualte; lateral leaflets
similar to terminal one. Racemes ebracteate; fruiting pedicels erect to ascending, 7–22 mm.
Sepals oblong, 2–3.5 × 2–2.5 mm, erect, base of lateral pair slightly saccate; petals white,
broadly obovate, 7–10 × 3–6 mm, clawed, apex rounded; median filament pairs 3–4 mm, lateral
pair 2–3 mm; anthers oblong, 1–1.5 mm. Fruit linear, 1.6–4 cm × 1–1.3 mm; valves glabrous;
style 0.7–3 mm; ovules and seeds 14–24 per fruit. Seeds brown, oblong, ca. 1.5 × 1 mm. 2n = 28,
42.
Flowering: Jul–Aug.
Habitat: moist streamsides, meadows, river gravel, mesic grounds, wet tndra, moist humus, scree
slopes, calcareous fellfield.
Elevation: 50–950 m.
Distribution: Russia (Far East), United States (W Alaska).
11
Cardamine bodinieri (H. Léveillé) Lauener, Notes Roy. Bot. Gard. Edinburgh 26: 336. 1965;
Dentaria bodinieri H. Léveillé, Repert. Sp. Nov. Regni Veg. 8: 452. 1910. TYPE: China,
Guizhou, Kouy-Yang, May 1910, Bodinier s.n. (holotype, E).
Herbs, perennial. Rhizomes present. Stems ca. 60 cm, erect, simple or branched above,
glabrous except for leaves. Basal leaves withered by flowering; cauline leaves 5−8, simple,
deeply 3-lobed; petiole 0.5−5 cm, with a broad wing 2−6 mm wide extending along entire length,
base amplexicaul; auricles broadly oblong to ovate, to 6 × 4 mm; terminal leaf lobe ovate to
ovate-lanceolate, 4−8 × 2−4 cm, sparsely hirsute with stout trichomes to 0.4 mm, sessile and base
decurrent with lateral lobes, margin serrate-crenate, ciliolate with trichomes less than 0.1 mm,
teeth mucronate, apex acute to acuminate; lateral lobes similar to terminal but smaller. Raceme
ebracteate, several flowered; fruiting pedicels divaricate-ascending, 5−10 mm, straight, glabrous.
Sepals ovate-oblong, 2−3 × ca. 1.5 mm, glabrous, lateral pair subsaccate; petals purple, obovate,
ca. 7 × 3 mm, not clawed; median filaments pairs 2.5−3 mm, lateral pair 1.5−2 mm; anthers ca.
0.6 mm; ovules 8−12 per ovary. Fruit linear, 0.8−1.2 cm × 1−1.2 mm; gynophore ca. 0.5 mm;
valves glabrous, smooth; style 0.5−2 mm. Seeds dark brown, oblong, 1.3−1.7 × 0.7−0.9 mm,
wingless. Flowering May−Jul.
Elevation: ca. 1100 m.
Distribution: China (Guizhou).
Cardamine bonariensis Persoon, Syn. Pl. 2: 195 (1807); C. flaccida subsp. bonariensis
(Persoon) O. E. Schulz. TYPE: Argentina, Buenos Aires, Commerson s.n. (holotype, P-JU;
isotype, P!).
Cardamine flaccida Chamisso & Schlechtendall, Linnaea 1: 21. 1826; C. bonariensis var.
flaccida (Chamisso & Schlechtendal) J. F. Macbride, Field Mus. Pub. Bot. 13, pt. 2: 962.
1938 TYPE: Chile. Talcaguano, 1816, Chamisso & Eschscholz s.n. (holotype, LE; isotype, B).
Nasturtium radicans Walpers, Nov. Act. Acad. Case. Leop.-Carol. 19(1). Suppl. 247. 1843;
Cardamine radicans (Walpers) Kuntze, Revis. Gen. Pl. 1: 21. 1891. TYPE: Chile, Valparaiso,
Collector ? (holotype,…).
Nasturtium turfosum Kunze ex Walpers, Nov. Act. Acad. Case. Leop.-Carol. 19(1). Suppl. 247.
1843. TYPE: Chile, Valparaiso, Pöppig 165 (holotype, ?; isotypes, G, ….).
Cardamine laxa Bentham, Pl. Hartweg. 158. 1845; C. flaccida prol. laxa (Bentham) O. E.
Schulz, Bot. Jahrb. Syst. 32: 448. 1903. TYPE: Ecuador, Popayan: near Tambo de Garbiel
Lopez in Páramo de Guanaco, Hartweg 880 (holotype, K!; isotypes, 2G!).
Cardamine nasturtioides Bertero ex Barnéoud in Gay, Fl. Chile 1: 113. 1846. TYPE: Chile,
Valparaiso, Berro, 1830, Bertero 174 (holotype, P!; isotypes, G!, P!; fragments, BAA!).
Cardamine laxa var. pumila A. Gray, Bot. U. S. Expl. Exped. Wilkes 1: 50. 1854. TYPE: Peru.
Culani to Casa Cancha, high Andes, collector? (holotype, GH?, or US?).
Cardamine minima Steudel, Flora 39: 410. 1856; C. flaccida subsp. minima (Steudel) O. E.
Schulz, Bot. Jahrb. Syst. 32: 451. 1903; C. bonariensis var. minima (Steudel) J. F. Macbride,
Field Mus. Pub. Bot. 13, pt. 2: 962. 1938. TYPE: Peru. San Antonio, Lechler 1811 (holotype,
P!).
Cardamine pusilla Philippi, Linnaea 28: 665. 1856; C. alsophila Philippi var. pusilla (Philippi)
Reiche, Fl. Chile 1: 99. 1896; C. flaccida subsp. alsophila (Philippi) O. E. Schulz prol. pusilla
(Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 450. 1903. TYPE: Chile, Mina de plata Las
Arañas, Prov. Santiago, Philippi s.n. (holotype, SGO-49308!; ?isotype, B!; fragments, BAA!).
12
Cardamine ramosissima Steudel, Flora 39: 409. 1856. TYPE: Chile, Rancagua [as Tagua-Tagua
in the original description], 1829, Bertero 147 (holotype, P!; isotypes, W!). There are several
of Bertero’s collection # 147, and some were collected prior or post 1829, as well as from
Rancagua or other localities. However, none of those should be considered as part of the type
collection of C. ramosissima.
Cardamine nemophila Philippi, Linnaea 30: 186. 185. 1859−1860. TYPE: Chile, Puerto Montt,
Feb 1858, Philippi s.n. (holotype, SGO-49306!).
Cardamine demissa Triana & Planchon, Annal. Sci. Nat., Bot. ser. 4, 17: 60. 1862. TYPE:
Colombia, Tolima, Llanitos du pied de Loma, 1844, J. Goudot s.n. (holotype, P!).
Cardamine axillaris Weddell, Ann. Sci. Nat. ser. 5, 1: 290. 1864. TYPE: Bolivia, La Paz,
Larecaja, vicinity of Sorata, river Challasuyo, 2600−2800 m, 1857−1858, Mandon 904
(holotype, P!; isotypes, BM!, F!, GH!. NY!, W!). A specimen each at G, K, and W were
collected at altitudes (3600−3800 m) and a river bank (Lacaita) different from those of the
holotype and isotype. Although they carry the exact other information of the type, they are not
considered herein as isotypes. How about the specimen at BM?
Cardamine axillaris var. tucumanensis Grisebach, Abh. Königl. Ges. Wiss. Göttingen 19: 71.
1874; C. flaccida var. tucumanensis (Grisebach) O. E. Schulz, Bot. Jahrb. Syst. 32: 451. 1903.
TYPE: Argentina, Tucuman, rivulis pr. Siambon, 25/31 Mar 1872, Lorentz 743 (holotype,
GOET, not seen; isotype, CORD!).
Cardamine andicola Philippi, Anal. Mus. Nac. Bot. 2: 1. 1891. TYPE: Chile, Aminchas,
between Ascotan and Pica, c. 3000 m, Feb 1885, F. Philippi s.n. (holotype, SGO-49317!;
fragments, BAA!). The protologue gives an altitude of 3000 m, but the specimen above says
3800 m. Muñoz-Pizarro (1960) also listed SGO-63913 as type collection, but I have not seen
this number.
Cardamine alsophila Philippi, Anal. Univ. Chile 81: 73. 1892; C. flaccida subsp. alsophila
(Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 449. 1903. TYPE: Chile, “In glareosis rivuli de
Aculeo,” Dec 1886, Philippi s.n. (holotype, ?SGO-71631!; see Muñoz-Schick, 1973).
Cardamine tridens Philippi, Anal. Univ. Chile 81: 74. 1892; C. alsophila Philippi var. tridens
(Philippi) Reiche, Fl. Chile 1: 98. 1896. TYPE: Chile, Prov. Aconcagua, Jahuel, pr. San
Felipe, Aug. Brochers 2276 (holotype, SGO-71620!; fragments, BAA!). Muñoz-Pizarro
(1960) listed two other collections, SGO-63882 and 63910 (fragments, BAA!), that I have not
yet seen, but if these do not carry the exact data given in the protologue, then they are not part
of the type collection.
Cardamine caespitosa Philippi, Anal. Univ. Chile 81: 79. 1892. C. alsophila Philippi var.
caespitosa (Philippi) Reiche, Fl. Chile 1: 99. 1896. TYPE: Chile, “Valle de de Santa Gertrudis
ad radicem borealem vulcani de Chillan,” Oct 1889, Fridericus Puga s.n. (holotype, SGO49312!; fragments, BAA!). Muñoz-Pizarro (1960) also listed SGO-63910 as type collection,
but I have not seen that.
Cardamine bracteata Philippi, Anal. Univ. Chile 81: 85. 1892; C. alsophila Philippi var.
bracteata (Philippi) Reiche, Fl. Chile 1: 99. 1896; C. flaccida f. bracteata (Philippi) O. E.
Schulz, Bot. Jahrb. Syst. 32: 448. 1903. TYPE: Chile, Andes of Santiago, Philippi s.n.
(holotype, SGO-49313!).
Cardamine cymbalaria Chodat & Wilczek, Bull. Herb. Boiss. Ser. 2, 2: 189. 1902. TYPE:
Argentina, Las Juntas, entrance of Valle de l’Atuel, E. Wilczek 447 (holotype, G!).
Cardamine flaccida prol. Dep. ressa O. E. Schulz, Bot. Jahrb. Syst. 32: 448. 1903. TYPE: Chile,
Talcahuano, 1828, Pöppig s.n. (holotype, W!).
13
Cardamine flaccida var. pilosa O. E. Schulz, Bot. Jahrb. Syst. 32: 448. 1903. TYPE: Chile, Juan
Fernandez Is., Nov 1864, Philippi s.n. (lectotype, here designated, B!; isolectotypes, SOG!,
W!).
Cardamine flaccida var. ebracteata O. E. Schulz, Bot. Jahrb. Syst. 32: 452. 1903. TYPE:
Mexico, Sand Andres, 1849, Chrismar s.n. (lectotype, here designated, B).
Cardamine flaccida var. macrantha O. E. Schulz, Bot. Jahrb. Syst. 32: 452. 1903. TYPE:
Mexico, San Pedro and S. Pablo, 1839, C. Ehrenberg 215 (holotype, B).
Cardamine killipii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 341. 1928. TYPE.
Colombia, Santander, eastern Cordillera, edge of Páramo de las Vegas, 3300−3700 m, 20−21
Dec 1926, E. P. Killip & A. C. Smith 15596 (holotype, B!; isotypes, F!, GH!, K!, NY!).
Cardamine flaccida var. ravenii Rollins, Crucif. Continental N. Amer. 273. 1993. TYPE:
Mexico, Chiapas, Muncicpio de Tenejapa, Banbil, 9,100 ft, 10 Oct 1965, D. E. Breedlove &
P. H. Raven 12909 (holotype, GH!).
Cardamine flaccida var. turfosiorum Rollins, Crucif. Continental N. Amer. 273. 1993. TYPE:
Costa Rica, Cartago,: Cordiller de Talamanca, near the Pan American Highway, ca. 2,000 m,
25 Aug 1961, C. Weber 6065 (holotype, GH!).
Herbs, perennial, usually rooting from lower nodes, glabrous or sparsely pubescent.
Rhizomes present. Stems (3−)7−35(−50) cm, erect, decumbent, or prostrate, simple or branched
above and/or below. Cauline leaves (0.8−)2−8 cm, petiolate, not auriculate, pinnately compound
or trifoliolate, rarely simple; petiole (0.5−)1−4(−5.5) cm; terminal leaflet (or leaf blade in simple
leaves) 0.5−2.5 cm, orbicular, subreniform, ovate, oblong, or lanceolate, base cuneate, obtuse, to
cordate, margin entire to repand or coarsely dentate to lobed, apex rounded to acute or
subacuminate, petiolule to 2 cm; lateral leaflets 1−3(−4) on each side of rachis, similar to
terminal leaflet but smaller and often with shorter petiolules, base cuneate or somewhat oblique,
usually absent and leaves simple in submerged plants; uppermost leaves smaller, with fewer
leaflets or all simple. Racemes bracteate throughout or at least along proximal half, rarely only
lowermost few flowers bracteates, many flowered,; rachis straight or rarely slightly flexuous;
fruiting pedicels (0.5−)0.8−2(−2.6) cm, erect to ascending, slender, straight or curved. Sepals
1.4−2 × 0.5−0.8 mm, oblong, caducous; petals 2−4.5 × 0.7−1.5 mm, white, spatulate, apex
obtuse; filaments 1.5−2.5 mm; anthers oblong to ovate, 0.4−0.5 mm; ovules 20−40 per ovary.
Fruits (0.7−)1.5−2.2(−3) cm × 0.7−1.2(−1.5) mm; style 0.5−1(−1.8) mm. Seeds light brown,
oblong, 0.8−1 × 0.6−0.8 mm.
Flowering: all year.
Habitat: ponds, streams, seepage areas, bogs, moist slopes, wet bluffs, muddy banks, wet paramo,
moist turf, gravel, wet grass fields, swales, marshes, ponds.
Elevation: 0−4500 m
Distribution: Argentina (Buenos Aires, Catamarca, Chubut, Cordoba, Entre Rios, Jujuy, La
Rioja, Mendoza, Misiones, Neuquén, Salta, Santa Cruz, Tucumán), Bolivia (Cochabamba, La
Paz, Potosí, Santa Cruz, Tarija), Brazil (Paraná, Santa Catarina, Rio Grande do Sul), Chile
(Región I, IV, V, Santiago, VI, VII, VIII, IX, X, XII, Juan Fernandez Is), Colombia
(Antioquia, Boyacá, Caldas, Cauca, Cundinamarca, Huila, Nariño, Norte de Santadaer,
Putumayo, Quindio, Putomayo, Santador, Tolima, Valle), Costa Rica, Ecuador (Azuay,
Bolivar, Cañar, Carchi, Chimborazo, Cotopaxi, Imbabura, Loja, Morona-Santiago, Napo,
Pastaza, Pichincha, Tunguarahua, Zamora-Chinchipe), Guatemala, Paraguay, Peru
(Amazonas, Ancash, Apurimac, Ayacucho, Cajamarca, Cuzco, Huánuco, Junin, La Libertad,
Lima, Pasco, Puno), Uruguay (Canelones), Venezuela (Merida).
14
Specimens examined: ARGENTINA. Buenos Aires: San Isidro, Parodi 10536 (F), Boelcke
3306 (BAA); Isla Santiago, Cabrera 2883 (LP, SI), Cabrera 3366 (G, GH, LP, NY, SI); Punta
Lara, Pastore-Troncoso 981 (SI); Anchosena, Hunziker 6279 (CORD, SI); Tigre, Burkart 149
(SI); La Plata, Krapovickas 530 (BAA). Catamarca: Las Chacritas, Boelcke et al. 5525 (BAA);.
Dep. Ambato, Las Chacritas, 27°50′S, 65°55′W, Boelcke et al. (BAA); Andalaglá, Esq. Grande,
Jörgensen 1441 (BAF, GH, MO, US), Dep. Belén, Nacimiento de San Antonio, Cabrera et al.
24696 (LP, SI). Chubut: Dep. Futaleufú, lago Futalufguen, Correa et al. 4168 (BAA, BAB,
BACP). Cordoba: Punilla, Route 20, 7 km W Tanti, Marínez-Laborde & Bernardello 30
(BACP); Sierra Chica, Lossen 279 (MO). Dep. San Alberto: Pampa de Achala:, Estancia San
Miguel, 25 km S ruta 20, Hunziker & Moscone 25099 (CORD). Entre Rios: Concepcion del
Uruguay, Burkart 17987 (BAA, SI); Concordia a Los Charrnas, Troncoso et al. 1994 (SI). Jujuy:
Valle Grande, arroyo de las Losetas, Cabrera et al. 27883 (MO, SI); Tumbaya, Ruta 52, Abra de
Potrerillos, 25 km de Purmamarca camino a Susques, 24°40′S, 65°37′W, Morrone et al. 2741
(SI). Dep. Tilcara: Casa Colorado, La Aguidita, Cabrera & Hernandez 13947 (LP). La Rioja.
Dep. Famatia/Chilecito: La Hoada, Kurtz 15105 (CORD); Sierra de Famatina, La Mesada,
Sparre 8885 (LIL). Mendoza: Dep. Malargüe, Invernada del Viejo, 35°43′S, 70°10′W, Prina et
al. 2196 (SI, SRFA). Misiones: Dep. Candelareia: Bompland, Nov. 1910, Hansen s.n. (BAB).
Dep. Guaraní, Predio Guarani, Tressens et al. 6417 (GH, NY). Neuquén: Dep. Minas, Las
Ovejas, 37°01′S, 70°45′W, Boelcke et al. 13551 (BAA, BAB, BACP, LIL, SI); Valle del Pichi
Neuquén, Boelcke et al. 13642 (BAA, BACP); confluencia del río Varvarco con el arroyo
Turbio, Boelcke et al. 14361 (BAA, BAB, BACP, SI); Cajón de los Chenques, 36°28′S,
70°48′W, Boeclcke et al. 13878 (BAA, BAB, BACP, SI); Aduana Vieja, 36°50′S, 71°04′W,
Boelcke et al. 10830 (BAA, BAB, SI). Salta: Cachi, rout du Cerro Negro, Km 20.5, Charpin &
Lazare 24040 (G); Cafayate, Krapovickas & Cristóbal 20682 (BAA, LIL). Dep. Capital: Salta,
Nov. 1892, Kuntze s.n. (CORD). Dep. Guachipas: Pampa Grande, Kurtz 12646 (CORD). Dep.
Santa Victoria: camino de Los Toldos a Huaico Chico, Legname & Cuezzo 9618c (LIL). Santa
Cruz: Dep. Río Chico, Río Oro, 47°26′S, 72°05′W, Boelcke et al. 12885 (BAA, BAB, SI).
Tucumán: Alpachiri, Río Cochuna, Burkart 26540 (SI, US). Dep. Tafí: Quebrada Honda,
Sparre 9325(LIL); Tafi del Valle, Diers 75 (SI). Dep. t Chicligasta, arroyo Celeste, R 65, Km 28,
27°20′S, 65°50′W, Boelcke et al. 5468 (BAA), Boelcke et al. 5465 (BAA). BOLIVIA. Unduavi,
Rusby 1366 (NY, US); Bermejo, Fiebrig 2086 (G), Fiebrig 2184 (G). Cochabamba: Cercado,
Cerro Multi Mayu, 17°21′S, 66°06′W, Ritter 1891 (MO); Chapare, Río Chaqui Mayu, nr hway,
Ritter & Crow 1102 (MO); Chapare, Jutun Corral, Río Candelaria, 17°16′S, 66°55′W, Ritter &
Crow 2160 (MO); Sacaba, Incachaca, Steinbach 5744 (K, SI), Steinbach 5949B (SI). La Paz:
Weddell s.n. (P); Murillo, 11.4 km E Ventilla on rd to Lambate, 16°34′S, 67°54′W, Solomon
15163 (GH, MO); Murillo, Asplund 597 (UPS), Asplund 612 (UPS); Ingavi, Huacullani, Beck
1036 (GH, LPB, MO). Potosí: Cerranía del Khare-Kare, Laguna Chalaviri, Schulte 57 (MO).
Santa Cruz: Yungas de San mateo, Steinbach 8522 (GH, K, LIL, NY). Tarija: entre Tarija y
Narvaes, Zuloaga et al. 1289 (SI). BRAZIL: Paraná: Calmon, Dusén 9298 (S). Santa
Catarina: Serra do Oratório, Reitz & Klein 6985 (US); Estrada Dona Francisca, Joinvile, Reitz &
Klein 4661 (US), Mun. Bom Retiro, Campo dos Padres, Smith, Reitz & Klein 7687 (US);
Pinheiral, Riozinho, Smith & Kline 7911 (GH). Rio Grande do Sul: São Francisco de PaulaLajeado Grande, Wasum 3413 (G). CHILE. I: Parinacota, Salar de Surire, 18°47′S, 69°06′W,
Argomedo 47 (CONC); Camino entre Putre y Pacollo, 18°47′S, 69°31′W, Arroyo 84-882
(CONC). IV: Coquimbo, Illapel, La Vega Escondida, E Cuncumen, Morrison 16954 (DS, UC,
US); Limarí, Cordillera de Ovalle, Río Molles, 30°43′S, 70°31′W, Jiles 3695 (CONC); Chopa,
15
Illapel, Fundo El Mollar-Los Vilos, 31°48′S, 71°29′W, Schlegel 3809 (CONC); Dep. Elqui,
Quebrada Las Trancas, Km 26 del camino Vicuña a Hurtado, Wagenknecht 4095 (SI). V: Río
Colorado, Los Andes, [32°53′S, 70°25′W], 23 Sept. 1951, Salazar s.n. (CONC); Cerro de la
Campana, Cajón Grande, cerca de la Poza del Coipo, [32°57′S, 71°08′W], Garaventa 4618
(BACP, CONC, SI), Garaventa 4616 (BACP, CONC, SI); Estero de Limache, [33°01′S,
71°16′W], Garaventa 2862 (BAA, BACP, CONC, SI), Garaventa 2687 (BAA, SI); Laguna
Verde, [33°03′S, 71°42′W], Garaventa 2396 (BAA, BACP, CONC, SI), Garaventa 2858 (SI);
Portezuelo de San Pedro, Limache, [32°57′S, 71°08′W], Garaventa 3380 (CONC), Garaventa
2208 (BAA, BACP, SI); Las Zablas, [33°21′S, 71°34′W], Garaventa 2719 (BAA, CONC);
Quintero, [32°46′S, 71°31′W], Richter 19636 (CONC), Gunckel 23792 (CONC); Quillota,
[32°53′S, 71°15′W], Oct. 1969, Martinez s.n. (CONC); Laguna de Campiche, 32°46′S, 71°32′W,
Gunckel 39475 (CONC); Fundo Perales, [33°05′S, 71°36′W], Sept. 1912, Jaffuel s.n. (CONC);
Valle de Oca, [32°50′S, 71°07′W], Schlegel 1162 (CONC); Valpariso, Claude-Joseph 3630
(US); Catapilco, Philippi 62a (SGO); Papudo, Zöllner 8306 (MO, NA, NY); Hacienda El Pangal,
near Limache, Killip & Pisano 39744 (US); Chapparro, Garaventa 2721 (BAA, BACP, SI).
Santiago: Potrero Grande de las Condes, [33°26′S, 70°20′W], 3 Jan. 1936, Grandjot s.n.
(CONC); Peñalolén, [33°28′S, 70°32′W], Gunckel 23310 (CONC); Peñalolén, 1 Nov 1881,
Philippi s.n. (SGO), Ortuzar Montt 202 (CONC); between P. Caldera and Maitenes, [33°16′S,
70°21′W], Skottsberg & Sparre 11097 (CONC); Santuario de la Naturaleza Yerba Loca, 2.5 km
from Villa Paulina, Sweeney 266 (MO); Til-Til, Looser 3352 (SI); Puente Alto, Garaventa 1711
(BAA, BACP, SI). VI: Colchagua, 1842, Cuming s.n. (G, W). VII: Cordillera de El Planchón,
[35°10′S, 70°35′W, Barros 2783 (CONC, SI); Liico, Curicó, [34°46′S, 72°041′W], Barros 2796
(CONC); Itahue, Fundo “El Colorado,” a la orilla del Río Claro, [35°08′S, 71°21′W, Garaventa
4515 (BACP, CONC, SI); Cerro de Talca, [35°36′S, 71°12′W], 10 Jan. 1915, Silva s.n. (CONC);
El Picazo, Talco, [35°32′S, 71°0′W], Barros 2800 (CONC, SI); Predio Venecia, I km al sur de
Talca, 35°27′S, 71°39′W, Rodríques & Baeza 2401 (CONC). VIII: San Rosendo, ClaudeJoseph 4382 (CONC, US); Cordillera de Nahuelbuta, cerro Pino Huacho, 37°41′S, 73°14′W,
Marticorena et al. 1581 (CONC); Sta. Bárbara-Fundo Ancud, [37°42′S, 71°58′W], Pfister 21312
(CONC), Pfister 664 (CONC); Gualpen, [36°47′S, 73°09′W], Barros 1217 (CONC); La Toma,
[35°50′S, 73°02′W], Junge 2961 (CONC); Camino Concepción a Coronel, Km 15.5, 36°53′S,
73°08′W, Parra & Rodríguez 32 (CONC); Arauco, camino desde Cañete a Contulmo, 1 km antes
del fundo La Passión, [37°53′S, 73°20′W], Baeza & Kottirsch 1436 (CONC). IX: Fundo Solano,
Los Alpes, Cordillera de Nahuelbuta, Eyerdam 10214 (F, UC, US); Contulmo, Cerro Santa
Elena, [38°0′S, 73°148′W], Ricardi 9239 (CONC); Saboya, [38°0′S, 72°48′W], Montero 5348
(CONC, SI); Cautin, Dep. Victoria, road 14 km from Termas de Tolhuaca to Laguna de Malleco,
Morrison & Wagenknecht 17467 (UC). X: Valdivia, Sept. 1895, Buchtien s.n. (F, GH, US); Río
Valdivia, [39°50′S, 73°14′W], Gunckel 37606 (CONC); San Juan, F. Philippi 60a (SGO);
Llanguihue, Los Santos, [44°08′S, 72°25′W], Villagrán 67663 (CONC); Riñihue, [39°50′S,
72°25′W], Gunckel 5860 (CONC), Santesson 1106 (S); Estancilla, [39°57′S, 73°19′W], Junge
2564 (CONC); Cumleufu (Trumao), [40°21′S, 73°10′W], Hollermayer 637a (CONC); La
Aguada, [39°53′S, 73°25′W], Gunckel 2778 (CONC, GH). Dep. Castro, Lago Huillinco,
42°42′S, 73°50′W, Marticorena et al. 143 (CONC). XII: Isla Long., 52°18′S, 73°38′W,
16
anonymous s.n. (SGO-63899). Juan Fernandez Is.: Falls V. Paugal, 4 April 1920, Bryan s.n.
(RSA); Masatierra, [33°380′S, 78°52′W], Sparre 61 (CONC), Gunckel 72958 (CONC),
Skottsberg 61 (NY, US), Skottsberg 219 (NY, P, US), Skottsberg 603 (US), Meyer 9589 (GH);
Mas Atiera, Pangal Falls, Meyer 9589 (CONC). COLOMBIA. Antioquia: San José de San
Andrés, Correa & Velásquez 53 (US). Boyacá: Sierra Nevada del Cocuy, El Llano de
Deslenguadas, Finca Ritacuva, Barclay & Juajibioy 7248 (GH, MO); Quebrada de San Paulino,
El Morrón, Cuatrecasas 1380 (F); Sierra Nevada del Cocuy, above Guican, Grubb et al. 86 (K,
US). Caldas: rd to Nevado, between rd to Libano and Sta. 10, Barclay & Juajibioy 6401 (GH,
MO); Cordillera Central, Paramo del Nuevado del Ruiz, Humbert et al. 27044 (P, SI), 27006 (P);
Nevado de Santa Isabel, páramos de la Laguna del Mosquito, Cuatrecasas 23242 (US); Páramo
del Quindio, Pennell & Hazen 9974 (NY). Cauca: Puracé, Parque Nacional Natural de Puracé,
Lozano et al. 4542 (F, NY); Puracé, Sneidern 2274 (GH, NY, S); Cabeceras del Río Páez,
Laguna del Páez, Cuatrecasas 19078 (GH, K, US); P’aramo del Puracé, San Francisco,
Cuatrecasas 14567 (US); Macizo Colombiano, alrededores de La Hoyala, Idrobo, Pinto &
Bischler 3548 (P); Mt Purace, Rio Anambiu, Killip 6771 (GH, NY). Cundinamarca: Macizo de
Bogotá, Quebrada del Rosal, Cuatrecasas 5702 (F); Macizo de Bogotá: Quebrada de San
Cristóbal, Cuatrecasas 5121 (F); Old Spanish rd from San Cristóbal to Ubaque, Finca El Delirio,
Schiefer 911 (GH, MO, NY); Mt Chuscal, W Zipaquira, Pennell 2603 (NY). Huila: Balsillas,
Rusby & Pennell 715 (GH, MO, NY); Rio Villalobos, vicinity or Río Suazita, Schultes &
Villarreal 5613 (GH, NY). Nariño: Mariña, near village Chiles, E Volcán Chiles, on the
Colombia-Ecuador border, Wiggins 10585A (DS). Norte de Santadaer: 23 km N Toledo up Río
Culaga, Fassett 25987 (US); NE of La Belleja, Fassett 25812 (GH). Putumayo: 5 km S of
Sibundoy, Chindoy 96 (GH) Quindio: Mpo Salento, Vereda Cocóra, 4°40-45′N, 75°20-25′W,
Luteyn, Fuertes & Rangel 13073 (GH, MO, NY). Putomayo: Pato-Macoa rd, near Mirador,
Croat 51786 (MO); 5 km S of Sibundoy, Bristol 434 (GH). Santador: between La Paz and
Velez, Fassett 25141 (GH). Tolima: Rosalito, near Páramo de Ruiz, Pennell 2976 (GH, K, MO,
NY). Valle: cabecerass del r’io Tuluá, Quebrada de Las Vegas, Cuatrecasas 20259 (P, US),
Cuatrecasas 20392 (P, US). ECUADOR. Vicinity of Huigra, Rose & Rose 22227 (NY). Azuay:
road Gualaceo-Limón, km 25.2, 3°0′29″-51″S, 78°39′43″-78°40′2″W, Jørgensen et al. 1882
(MO, NY). Bolivar: Hacienda Talahua, Penland & Summers 638 (F, GH). Bioco: Malabo-Pico
Basilé, km 3, Carvalho 2154 (MO). Cañar: Ingapirca, 2°30′S, 78°53′W, Kohn 1460 (MO); near
El Tambo, Camp E-3984 (GH, NY). Carchi: between Tulcan and Maldonado, 0°47′N, 77°58′W,
Eriksen 59005 (AAU, MO, NY, QCA); 28 km N El angel, 0°43′N, 77°47′W, Holm-Nielsen
23884 (AAU, NY). Chimborazo: Nevado El Altar, near Río Blanco, 1°40′S, 78°30′W, Juncosa
848 (GH, MO); Riobamba Canton, Parroquia Pungala, Comunidad Alao-Lactapamba, Parque
Nacional Sangay, Páramo de Culebrillas, 1°58′S, 78°28′W, Cerón, Montesdeoca & Craing
11806 (MO, QCNE); Urbina Pass, Nudo de Igualata, Penland & Summers 455 (F, GH); Páramo
de Carnicería, Bolcán Sangay, Acosta Solís 7679 (F); Chimborazo/Morona Santiago, Cerros
Yuibug, Pailacajas, 1°45′S, 78°27′W, Sklenar & Sklenarova 3084 (MO, PRC). Cotopaxi: Muro
Urcu, 0°47′S, 78°24′W, Holm-Nielsen 16480 (AAU, MO, NY, QCA); Parque Nac. de Cotopaxi,
Quebrada Mishaguaico along road to Limpio Pungo, 0°38′S, 78°28′W, Laegaard 101467 (AAU,
MO); Río Chalupas, at Cotopaxi/Napo, 0°50′S, 78°21′W, Laegaard 101739 (AAU, MO).
Imbabura: Lake Cuicocha, Isolte Chica, Asplund 7159 (GH, K, LIL, NY, P, S); NE side of
Cayambe Mt, Cazalet & Pennington 5776 (B, K, NY, UC, US); km 28-31 Yahuarcocha-Mariano
Acosta, 0°20′N, 77°58′W, Laegaard 101216 (AAU, MO); NW slopes of Cayambe, Wiggins
10374 (DS, GH). Loja: Loja-Zamora rd, km 15, 3°58′S, 79°08′W, Madsen 74028 (AAU); 12 km
17
on Loja-Zamora rd, 3°55′S, 79°09′W, Ellemann 75382 (AAU, MO); 12 km NW Saraguro on
Loma Paredones, 3°36′S, 79°10′W, Ellemann 91666 (AAU, MO). Morona-Santiago: between
Cebadas and Zuñac on Riobama-Macas rd, 2°12′S, 78°23′W, Croat et al. 86242 (MO);
Gualaceo-Limón rd, 0.6 km E of Azuay border, 28.5 km E Gualaceo, 3°00′S, 78°42′W, Croat &
Menke 89183 (MO); Parque Nacional Sangay, between Yanayacu and Culebrillas, 2°0′S,
78°30′W, Clark et al. 1867 (MO); between Mendez and Paute, 25 km W Méndez, 2°38′S,
78°25′W, Croat et al. 90779 (MO). Napo: Laguan Papallacta, 0°23′S, 78°9′W, Laegaard
103120 (AAU, MO, QCA); Laguna San Marcos, 0°7′S, 77°58′W, Øllgaard et al. 34054 (AAU,
MO, NY, QCA); 6 km NE of km 45 on Salcedo-Napo rd, 0°56′S, 78°23′W, Laegaard 53365
(AAU, MO, QCA); NW slope of Antisana, N Lago Mauca-Machay, 0°26′S, 78°9′W, HolmNielsen 20751 (AAU, MO, NY, QCA); Quijos Cantón, Laguna La Mica, 0°32′S, 78°13′W,
Vargas & Narváez 2150 (MO); Río Aliso, 8 km SE Cosanga, 0°36′S, 77°56′W, Vargas et al.
2934 (MO); Baeza, 0°28′S, 77°53′W, Balslev & Madsen 10301 (F); Laguna Encantada, 1°11′S,
78°12′W, Holm-Nielsen 41979 (AAU, NY). Napo-Pastaza: flat below Ainchilibi, Barclay &
Juajibioy 9056 (MO); Baeza, Gilli 354 (W). Pichincha: Tambillo, Asplund 6413 (K, NY);
Carretera antigua Chillagallo-San Juan-Chiriboga-Empalme, km 27, 0°17′S, 78°38′W, Zak 862
(AAU, K, MO, NY); Pifo-Pintag, 0°19′S, 78°17′W, Laegaard 102255 (AAU, MO, QCA); Rucu
Pichinca, 0°10′S, 78°34′W, Sklenar & Kostechova 249 (MO, PRC); Oyacachi, Acost-Solis 11102
(F) Tunguarahua. Santiago de Pillaro, Parque Nacional Llanganates, Valle de los Frailejones,
1°10′S, 78°15′W, Narváez et al. 494 (MO); Mocha, Parroquia Pinguili, 2 km aantes de la Base
del Lado Oriental del Caryguayrazo, Cerón 19261 (MO); Banos Cantón, Parque Nacional
Llanganates, 0°10′S, 78°15′W, Vargas, Ronquillo & Granda 2742 (MO); 6 km S Mocha, half
way between Ambato and Riobamba, 1°27′S, 78°41′W, Iltis & Iltis E-453 (MO); between Leito
and la Cima, Acosta-Solís 9064 (F). Zamora-Chinchipe: km 17 on Loja-Zamora rd, 3°59′S,
79°8′W, Holm-Nielsen & Jeppesen 3546 (AAU, B, MO, S); 13 Km E pass on Loja-Zamora rd,
3°58′S, 79°05′W, Øllgaard et al. 90892 (AAU); parque nacional Podocarpus, YanganaValladolid rd, km 26, 4°29′S, 79°9′W, Madsen et al. 75769 (AAU). PARAGUAY. Cordillera de
Villa Rica, Balansa 2622 (P); 81.3 km from border of Loja Prov. on road from Zamora to Loja,
4°08′S, 79°56′W, Croat & Mencke 89834 (MO). PERU. Pueblo, 3 miles below Ambo,
Tomaiquichua, Macbride & Featherstone 2431 (F, G, NY). Amazonas: Chachapoyas,
Leimebamba-La Joya trail, Boeke 1857 (SI). Ancash: Cajatambo, Weberbauer 2688 (G);
Huaylas, Quebrada Alpamayo, 8°50′S, 77°42′W, Smith et al. 9836 (MO); Huaraz, Quebrada
Shallap, 9°30′S, 77°23′W, Smith et al. 9685 (MO); Yungay, Quebrada Ranicuray, 8°59′S,
77°34′W, Smith et al. 10413 (GH, MO); Carhuaz, Quebrada Ishinca, 9°22′S, 77°25′W, Smith et
al. 9542 (GH, MO). Prov. Bolognesi, Racrachaca, Cerrate 711 (GH). Apurimac: 4 km E of
Cincacasa at km 177 from Puquio on rd to Chalhuanca, Iltis et al. 522 (GH). Ayacucho: Prov.
Lucanas, 8 km NW Puquio, above San Juan, Iltis et al. 468 (GH). Cajamarca: San Miguel,
Piedra Angosta, Llatas 809 (MO); Cajamarca, Pampa Larga, N Minera Yanacocha, Sánchez
7144 (F); Contumazá, Jalca de las Quinuas, Sagástegui et al. 10093 (MO, NY); Celendín,
Sendamal, Sagástegui et al. 12095 (MO, NY). Cuzco: Pancartambo, Hac. Churu, Herrera 2331
(F); La Raya, Pennell 13498 (F); Paucartambo to Tres Cruces, Cerro de Cusilluyoc, Pennell
14164 (F, GH, NY). Huánuco: Huánco, Carpish pass, Asplund 12814 (K, UPS); Carpish pass,
84 km from Tingo María on hwy to Lima, Allard 20992 (US); between Huico and Tingo Maria
18
in valley of Río Huallaga, near km 443.5, 9°38′S, 76°03′W, Croat 57830 (GH, MO); 13 km S
Chinchao, Gentry et al. 19270 (GH, MO, NY); 46.8 km NE Huánco on rd to Tingo María,
Plowman & Rury 11112 (F). Junin: Yauli-San Cristobol road, ca. 8 km from Yauli, Duncan et
al. 2663 (MO); km 125 on route 20A from Lima to La Oroya, Davidson 3346 (MO). La
Libertad: near Trujillo, Morrison 8879 (UC). Lambayeque: Lambayeque, El Potrero, Penachi,
Llatas 1427 (F); Ferrenafe, ca. 4 km NW Incahuasi, below Cerro Punamachay on trail to Laguna
Hualtaco, Dillon & Skillman 4161 (F). Lima: Río Blanco, Killip & Smith 21712 (F, NY); Prov.
Yuayos, Cruz-pampa, Cerrate 1113 (GH). Pasco: Oxapampa, 2-4 km N Mallampampa, 10°32′S,
75°45′W, Smith & Canne 5850 (MO). Puno: N of Puno at km 4 on road to Juliaca, Hutchison
1821 (MO, UC); Occa Pampa, Shepard 92 (GH). URUGUAY. Canelones: Pando, Herter 2258
(MO). VENEZUELA. Merida: vicinity of Timotes, Pittier 12671 (G, NY); Páramo de
Mucubají, alrededores de La Laguna Grande, Barclay & Juajibioy 9629 (MO); 15 km WSW
Santo Domingo or road to Merida, 8°48′N, 70°46′W, Liesner 13767 (MO); Miranda, Quebrada
El Turmero (afluente del Río Motatán), 10.4 km de El Aquila por la carretera a Pnñango, Berry
& Calvo 4380 (MO); slightly below La Laguna Negra on trail to Laguna de Mucubaji, Barclay &
Juajibioy 9803 (GH, MO); Rangel, Páramo de Motumbo, along border of divide to Laguna Las
Parias, Stergios, Dorr & Wurdack 20519 (MO, US); Laguna Mucubají, above Los Apartaderos,
Steyermark 57498 (F, GH), Vareschi 2123 (M)
Undoubtedly, Cardamine bonariensis is the most variable Central and South America species
of the genus, and it exhibits tremendous altitudinal and latitudinal ranges accompanied by a wide
variation of water bodies in which it frequently grows. It occupies altitudes from sea level (Parra
& Rodríguez 32 at CONC) to 4500 m (Schulte 57 and Duncan et al. 2663, both at MO) and
diverse habitats ranging from somewhat dry or slightly mesic areas all the way to completely
submerged forms. This ecological diversity goes hand in hand with tremendous morphological
variation and phenotypic plasticity. Perhaps the most variable parts are the leaves and they range
from compound and 3−9-foliolate terrestrial forms to those with exclusively simple leaves
growing in aquatic or mesic habitats with saturated soils. The leaf and leaflet margin also varies,
and in the aquatic forms it is always entire or repand whereas in the terrestrial forms they are
dentate or lobed. The presence of indumentum always seems to be found in the terrestrial forms.
The bracteate portion of the raceme also varies considerably, and some forms have racemes
bracteate throughout while others have only the proximal half is bracteate, and yet in other very
rare forms only the lowermost one or two flowers are bracteate. As for fruit length and
orientation, it is quite variable, and none of the morphological extremes in leaf morphology show
any correlation with fruit orientation and size. The morphological continuity between the various
forms often occurs within a given area, and the variation in morphology does not correspond
with geography. What is most urgently needed is to conduct extensive experimentation on plants
with identical genotypes grown under different ecological conditions. Because the plants are
rhizomatous perennial, one can easily conduct that type of experimentation. However, it appears
likely that some ecological differentiation would be found in such a variable species with
continuous variation from Mexico southward into Patagonia.
Schulz (1903), Boelcke & Romanczuk (1984), and Rollins (1993) adopted the name
Cardamine flaccida for the species, but the earliest name is C. bonariensis, and the types of the
two taxa are hardly distinguishable. However, Schulz (1936) recognized both C. bonariensis, C.
flaccida, and C. killipii as three related species. Boelcke (1967) recognized both C. bonariensis
and C. flaccida and separated C. bonariensis by having prostrate (vs. ascending or erect) stems,
flowers from axils of middle and upper leaves (vs. bracteate racemes), and variously oriented (vs.
erect) fruits. In my opinion, this complex represents a continuum the three species recognized by
19
Schulz (1936) of which represent only a fraction of the many extremes variously recognized as
species, subspecies, varieties, or forms. The type of C. bonariensis is a robust plant bracteate
throughout and with obovate, three lobed, simple, long-petiolate bracts and trifoliolate uppermost
cauline leaves. The basal portion of the plant is missing. The types of other extreme (initially
described as C. minima,C. demissa, C. pusilla, C. andicola, and C. axillaris) have simple,
orbicular to reniform, entire, short-petiolate leaves and bracts and rooting nodes. This form is
typical of the floating, submersed, or boggy plants, and often it forms dense mats. Its much
coarser form was recognized by Schulz (1928, 1936) and Macbride (1938) as C. killipii. The
other extreme, represented by C. nasturioides and C. alsophila, are erect plants of wet habitats
and have 7−9-foliolate cauline leaves, 3−5-foliolate bracts, and dentate to crenate leaflets. The
type of C. flaccida falls morphologically between those of C. bonariensis and C. nasturtioides.
There is no perfect solution that handles all of the tremendous variability to the satisfaction of
everyone. Schulz (1903), and to a limited extent Rollins (1993), took the position of according
infraspecific status to as many variants as they can and without consideration of the geographic
and morphological integrity of the variants. For examople, forms almost typical to the type of C.
killipii from Colombia and be found in Argentina (Parodi 10536 at F), Peru (Killip & Smith
21712 at F), Bolivia (Ritter & Garrett 1102, MO), and Ecuador (Sklenar & Sklenarova 3084 at
MO). Similar examples can also be cited for the other forms represented by the types of C.
bonariensis, C. nasturtioides, and C. minima. Indeed, Schulz (1903) left behind far more
additional variants that he did not name, and most of the taxa he recognized have sporadic
occurrence throughout South America similar to the example of C. killipii. Following and further
building on that approach, one would make the taxonomy of this complex far more confusing for
subsequent workers. The other extreme would be to recognize one, highly variable species
without infraspecific taxa and in the same way we handle the equally variable watercress,
Nasturtium officinale R.Brown. In the absence of extensive field and molecular studies on the C.
bonariensis complex, I am inclined not to recognize any infraspecific taxa.
Morrison 16954 is a mixed collection of Cardamine bonariensis and C. volckmannii, whereas
Holm-Nielson et al. 3546 is a mixed sample of C. bonariensis and C. ovata. Boelcke et al. 5465
(BAA) is a mixed collection of C. bonariensis and C. vulgaris.
Cardamine bonariensis is the most variable Central and South America species of the genus,
and it exhibits tremendous altitudinal and latitudinal ranges accompanied by a wide variation of
water bodies in which it frequently grows. Perhaps the most variable parts are the leaves and they
range from compound and 3−9-foliolate terrestrial forms to those with exclusively simple leaves
growing in aquatic or mesic habitats with saturated soils. The leaf and leaflet margin also varies,
and in the aquatic forms it is always entire or repand whereas in the terrestrial forms they are
dentate or lobed. The bracteate portion of the raceme also varies considerably, and some forms
have racemes bracteate throughout while others have only the proximal half is bracteate, and yet
in other very rare forms only the lowermost one or two flowers are bracteate. As for fruit length
and orientation, it is quite variable, and none of the morphological extremes in leaf morphology
show any correlation with fruit orientation and size. The morphological continuity between the
various forms often occurs within a given area, and the variation in morphology does not
correspond with geography. What is most urgently needed is to conduct extensive
experimentation on plants with identical genotypes grown under different ecological conditions.
Cardamine breweri S. Watson, Proc. Amer. Acad. Arts 10: 339. 1875. TYPE: United States,
California, near Sonora Pass, 8000–10,000 ft, 17 Jul 1863, W. H. Brewer 1890 (lectotype
designated by Detling (1937: 72), GH!).
Cardamine callosicrenata Piper, Bot. Gaz. 22: 488. 1896. TYPE: United States
20
Cardamine foliacea Greene, Pittonia 4: 201. 1900. TYPE: United States,
Cardamine hederaefolia Greene, Pittonia 4: 202. 1900. TYPE: United States, Oregon, E base of
Cascade Mts., 1893, Mrs. R. M. Austin s.n. (holotype, NDG).
Cardamine leibergii Holzinger, Contr. U.S. Natl. Herb. 3: 212. 1895; C. breweri var. leibergii
(Holzinger) C. L. Hitchcock, Univ. Wash. Publ. Biol. 17(2): 468. 1964; C. vallicola subsp.
leibergii (Holzinger) O. E. Schulz, Bot. Jahrb. Syst. 32: 523. 1903; C. sandbergii Holzinger,
Contr. U.S. Natl. Herb. 3: pl. 3. 1895. TYPE: United States, Idaho, Kootenai Co., near summit
of Packsaddle Peak, 6 Aug 1892, J. H. Sandberg, D. T. MacDougal, & A. A. Heller 856
(holotype,; isotypes, NY!; US!).
Cardamine modocensis Greene, Pittonia 4: 203. 1900. TYPE: United States, California, Modoc
Co., Lassen Creek, Aug. 1894, Mrs. R. M. Austin s.n. (lectotype designated by Jepson (Fl.
Calif. 2: 55. 1936), NDG).
Cardamine orbicularis Greene, Pittonia 4: 202. 1900; C. breweri var. orbicularis (Greene)
Detling, Amer. J. Bot. 24: 73. 1937. TYPE: United States,
Cardamine vallicola Greene, Pittonia 3: 116. 1896. TYPE: United States,
Herbs, perennial, glabrous or rarely sparsely pubescent near base. Rhizomes slender,
cylindrical, rarely slightly thickened at stem base, 1–3(–4) mm in diam.; stolons absent. Stems
(0.6–)1.5–6(–7) dm, erect or decumbent at base, simple or branched, glabrous or pubescent near
base. Rhizomal leaves absent; cauline leaves 3–8(–11), 3–5-foliolate or rarely only terminal
leaflet present, 2.5–12(–13.5) cm; petiole (0.7)1–4(–6) cm, base not auriculate; terminal leaflet
ovate to orbicular or rarely subcordate, 1.5–4(–5) × 1.5–3.5(–5) cm, petiolule 0.4–1.6 cm, base
truncate, rounded or cordate, margin crenate, dentate, sinuate, or to 11-lobed, puberulent or not;
lateral leaflets (when present) often considerably smaller and narrower than terminal one,
subsessile or on petiolule to 4 mm, margin same as terminal leaflet. Racemes ebracteate; fruiting
pedicels ascending to divaricate-ascending, (0.7–)1–2 cm. Sepals oblong, 2–3(–3.8) × 1–1.5 mm,
erect, base of lateral pair not saccate; petals white, oblanceolate, 3.5–6(–7) × 1.5–2.5(–3) mm,
not clawed, apex rounded or slightly emarginate; median filament pairs 2.5–3.5 mm, lateral pair
2–2.5 mm; anthers oblong, 0.7–1 mm. Fruit linear, 1.5–3.5 cm × 1–1.5 mm; valves glabrous;
style 0.2–1.5(–2.5) mm; ovules and seeds 14–28 per fruit. Seeds brown, oblong, 1–1.6 × 0.9–1.1
mm.
Flowering: Jun–Jul.
Habitat: streambanks, seepage, lake shores, creeks, wet meadows, swamps, ponds.
Elevation: 1200–3000 m.
Distribution: Canada (British Columbia), United States (E and N California, N Coloradoa, Idaho,
W Montana, Nevada, Oregon, N Utah, Washington, W and S Wyoming).
Cardamine bulbifera (Linnaeus) Crantz, Cl. Crucif. Emend. 127. 1769; Dentaria bulbifera
Linnaeus, Sp. Pl. 2: 653. 1753. TYPE: Herb. Burser XVIII(1): 82 (lectotype designated by
Marhold (2001: 45), UPS).
Cardamine bulbosa (Schreber ex Muhlenberg) B.S.P., Prelim. Cat. 4. 1888; Arabis bulbosa
Schreber ex Muhlenberg, Trans. Amer. Philos. Soc. 3: 174. 1793; Dracamine bulbosa
(Schreber ex Muhlenberg) Nieuwland, Amer. Midl. Naturalist 4: 40. 1915. TYPE:
Arabis rhomboidea Persoon, Syn. Pl. 2: 204. 1807; C. rhomboidea (Persoon) de Candolle, Syst.
Nat. 2: 246. 1821; Dentaria rhomboidea (Persoon) Greene, Pittonia 3: 124. 1896. TYPE :
Cardamine rhomboidea var. hirsuta O. E. Schulz, Bot. Jahrb. Syst. 32: 426. 1903. TYPE:
Cardamine rhomboidea var. parviflora O. E. Schulz, Bot. Jahrb. Syst. 32 : 426. 1903. TYPE:
Cardamine rhomboidea var. pilosa O. E. Schulz, Bot. Jahrb. Syst. 32: 426. 1903. TYPE:
21
Herbs, perennial, glabrous or sparsely pubescent distally. Rhizomes fleshy, tuberous at stem
base and sometimes at intervals, subglobose, lobed, 4–15 mm in diam.; stolons absent. Stems
(1–)2–6 dm, erect, simple, glabrous or sparsely pubescent on distal half with trichomes 0.02–0.1
mm. Rhizomal leaves simple, (2–)4–13(–16) cm; petiole (1.5–)2.5–10(–13) cm; blade reniform
to cordate or ovate, rarely oblong, (1–)2–4(–6) cm, base obtuse to cordate, margin repand or
entire, rarely shallowly dentate, not puberulent; cauline leaves (2–)4–10(–14), simple, not
auriculate at base, middle ones short petiolate, 3–6(–9) × 1–3(–4.5) cm, upper ones sessile, ovate
to oblong or oblong-linear to lanceolate, entire, repand, or dentate, minutely pubescent along
margin, Racemes ebracteate; fruiting pedicels ascending to divaricate, (1–)1.5–2.2(–3) cm.
Sepals oblong, 2.5–4.5 × 1.5–2 mm, erect, glabrous, base of lateral pair not saccate; petals white
or very rarely pale pink, obovate, (6–)7–12(–16) × 3–5 mm, short clawed, apex rounded; median
filament pairs 4.5–7 mm, lateral pair 2–3.5 mm; anthers oblong, 1–1.5 mm. Fruit linear, 2–3.5(–
4) cm × 1.4–1.7 mm; valves glabrous; style 2–4(–5) mm; ovules and seeds 14–24 per fruit. Seeds
dark orange to greenish yellow, oblong to circular, 1.7–2.1 × 1–1.4 mm. 2n = 16, 56, 64, 80, 96,
112.
Flowering: Mar–Jun.
Habitat: wet grounds, low woodlands, moss hummocks, alluvial woods, grassy floodplains, wet
pastures, meadows, pine lands, creek bottoms, stream banks, sandy bottoms, ditches, mesic or
wet forests, swamps, marshes, seepy bluffs.
Elevation: 0–850.
Distribution: Canada (Manitoba, Ontario, Quebec), United States (Alabama, Arkansas,
Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa,
Kansas, Kentucky, Louisiana, Maine, Maryland, Massachussets, Michigan, Minnesota,
Mississippi, Missouri, Nebraska, New Hampshire, New Jersey, New York, North Carolina,
Ohio, Oklahoma, Pennsylvania, South Carolina, E South Dakota, Texas, Tennessee, Vermont,
Virginia, West Virginia, Wisconsin).
Cardamine calcicola W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 11: 203. 1919. TYPE:
China, Yunnan, Mountains in the NE of the Yangtze bend, 27°45′N, 11,000–12,000 ft, Jul
1913, George Forrest 10471 (holotype, E!).
Herbs, perennial, sparsely to densely pilose on stem and petioles with crisped trichomes to
1.2 mm. Rhizomes thick, short, often with several stolons. Stems 10–35 cm, erect, simple, pilose
or distal parts subglabrous, not flexuous. Basal leaves rosulate, simple or rarely trifoliolate;
petiole 1–10 cm, often densely pilose; leaf blade or terminal leaflet reniform, 0.7–3 × 1–4 cm,
sparsely pilose or subglabrous, base cordate, margin subentire, repand, or 7–9-crenate, apex acute
or obtuse; cauline leaves 2–6, 3–7-foliolate; petiole 0.2–2.5 cm, not auriculate at base; terminal
leaflet obovate, 0.7–2.5 × 0.6–2.5 cm, with a petiolule 1–8 mm, sparsely pilose or subglabrous,
base cuneate and often decurrent with adjacent lateral leaflets, margin entire or obscurely to
strongly dentate, apex acute or rounded; lateral leaflets much smaller than terminal one, sessile or
rarely shortly petiolulate. Racemes ebracteate; fruiting pedicels divaricate or ascending, 0.7–1.6
cm, straight. Sepals oblong, 3–4 × 1–2 mm, margin and apex membranous, base not saccate;
petals white, obovate, 6–9 × 3–5 mm, apex rounded; median filament pairs 2.5–3.5 mm, lateral
pair 2–2.5 mm; anthers narrowly oblong, 1–1.3 mm; ovules 12–14 per ovary. Fruit linear, 2–3
cm × 1–1.5 mm; valves glabrous, smooth; style 0.5–2 mm. Seeds brown, oblong-ovate, 1.3–1.9 ×
0.8–1.3 mm, wingless.
Flowering: May–Jul.
Habitat: crevices of limestone cliffs, moist rocky pastures, valleys.
22
Elevation: 2600–3700 m.
Distribution: China (Yunnan).
Cardamine caldeirarum Guthnick
Cardamine californica (Nuttall) Greene, Fl. Francisc. 266. 1891; Dentaria californica Nuttall in
Torrey & A. Gray, Fl. N. Amer. 1: 88. 1838; D. integrifolia var. californica (Nuttall) Jepson,
Man. Fl. Pl. Calif. 426. 1925. TYPE:
Cardamine californica var. brevistyla O. E. Schulz, Bot. Jahrb. Syst. 32: 389. 1903. TYPE:
Cardamine californica var. fecunda O. E. Schulz, Bot. Jahrb. Syst. 32: 386. 1903. TYPE:
Cardamine californica var. pubescens O. E. Schulz, Bot. Jahrb. Syst. 32: 388. 1903. TYPE:
Cardamine californica var. robinsoniana O. E. Schulz, Bot. Jahrb. Syst. 32: 385. 1903. TYPE:
Cardamine cardiophylla Greene, Fl. Francisc. 266. 1891; C. californica prol. cardiophylla
(Greene) O. E. Schulz, Bot. Jahrb. Syst. 32: 388. 1903; C. californica var. cardiophylla
(Greene) Rollins, Harvard Pap. Bot. 4: 44. 1993; D. cardiophylla (Greene) B. L. Robinson in
A. Gray & S. Watson, Syn. Fl. N. Amer. 1(1): 155. 1895; D. californica var. cardiophylla
(Greene) Detling, Amer. J. Bot. 23: 576. 1936; D. integrifolia var. cardiophylla (Greene)
Jepson, Man. Fl. Pl. Calif. 426. 1925. TYPE:
Cardamine cuneata Greene, Bull. Calif. Acad. Sci. 1: 74. 1885; C. californica subsp. cuneata
(Greene) O. E. Schulz, Bot. Jahrb. Syst. 32: 386. 1903; C. californica var. cuneata (Greene)
Rollins; D. californica var. cuneata (Greene) Detling, Amer. J. Bot. 23: 576. 1936; D.
cuneata (Greene) Greene, Pittonia 3: 123. 1896. TYPE:
Cardamine sinuata Greene, Erythea 1: 148. 1893; C. californica var. sinuata (Greene) O. E.
Schulz, Bot. Jahrb. Syst. 32: 387. 1903; C. integrifolia var. sinuata (Greene) C. L. Hitchcock,
Univ. Wash. Publ. Biol. 17(2): 469. 1964; D. sinuata (Greene) Greene, Pittonia 3: 123. 1896;
D. californica var. sinuata (Greene) Detling, Amer. J. Bot. 23: 576. 1936. TYPE:
Cardamine pauscisecta Bentham, Pl. Hartweg. 297. 1848. TYPE:
Dentaria integrifolia Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 88. 1838; C. integrifolia
(Nuttall) Greene, Bull. Calif. Acad. Sci. 2: 389. 1887; C. californica prol. integrifolia
(Nuttall) O. E. Schulz, Bot. Jahrb. Syst. 32: 386. 1903; C. californica var. integrifolia
(Nuttall) Rollins, Harvard Pap. Bot. 4: 44. 1993; D. californica var. integrifolia (Nuttall)
Detling, Amer. J. Bot. 23: 586. 1936. TYPE:
D. integrifolia var. traceyi Jepson, Man. Fl. Pl. Calif. 426. 1925. TYPE:
Dentaria pachystigma S. Watson var. dissectifolia Detling, Amer. J. Bot. 23: 575. 1936; C.
pachystigma (S. Watson) Rollins var. dissectifolia (Detling) Rollins, Harvard Pap. Bot. 4: 45.
1993. TYPE: United States, California, Butte Co., Magalia, L. E. Detling 1631 (holotype,
DS!).
Herbs, perennial, glabrous or rarely minutely pubescent. Rhizomes fleshy, globose to ovoid
or suboblong, (3–)4–10 mm in diam., deeply underground; stolons absent. Stems (2–)2.7–6(–7)
dm, erect, simple, glabrous or rarely pubescent. Rhizomal leaves 3(or 5–7)-foliolate, or
occasionally simple, 8–25(–38) cm; petiole (5–)8–25(–32) cm; terminal leaflet or blade of simple
leaf ovate to orbicular to broadly cordate or reniform, (1.5–)2.5–7.5(–10) × (1.2–)2–9(–13) cm,
with a petiolule (0.7–)2–5(–11) cm, base obtuse to cordate, margin entire or dentate to shallowly
sinuate, often with apiculae at veins ending at margin, glabrous or minutely pubescent on veins;
lateral leaflets (when present) about as large as or smaller than terminal leaflet, petiolulate to
subsessile, margin same as terminal leaflet; cauline leaves 2–5, 3(or 5)-foliolate or rarely simple;
petiole 1–5(–9) cm, base not auriculate; terminal lobe broadly ovate to suborbicular or lanceolate,
rarely narrowly oblong, 1–7 × (0.5–)1–4.7(–6.5) cm, petiolulate or sessile; lateral leaflets similar
23
to terminal one but smaller, sessile, margin dentate or rarely entire. Racemes ebracteate; fruiting
pedicels ascending to divaricate, 1–3.3(–4.1) cm. Sepals oblong, 3.5–4.5(–5.5) × 1.5–2(–2.5)
mm, erect to ascending, base of lateral pair saccate; petals white to pale rose, often broadly
obovate, 8–13(–15) × 4–8 mm, clawed, apex rounded; median filament pairs 4–6.5 mm, lateral
pair 3–4 mm; anthers oblong, 1.2–1.7 mm. Fruit linear, 2.2–5.4(–6) cm × 2–3 mm; valves
glabrous; style 2–5(–6) mm; ovules and seeds 12–22 per fruit. Seeds dark brown, oblong to
broadly ovate, 1.7–2.8 × 1.2–1.8 mm. 2n = 32.
Flowering: Jan–May.
Habitat: wooded ravines, forest floor, shady slopes, open woods, shady rock crevices, stream
banks and bottoms, caynons, moist hillsides, cliffs.
Elevation: 0–1400 m.
Distribution: Mexico (Baja California), United States (California).
Cardamine calthifolia H. Léveillé, Bull. Acad. Géog. Bot. 24: 281. 1914. TYPE: China,
Yunnan, valley of Kiao-me-ti, 300 m, E. E. Maire s.n. (holotype, E!; isotypes, E!, P!).
Herbs, perennial, sparsely pilose at least on leaves. Rhizomes stout, to 7 cm in diam., with a
few stolons. Stems 11–30 cm, stout, thickest at base, erect, simple. Basal leaves fleshy, simple;
petiole (3.5)6–15(–20) cm, glabrous; leaf blade reniform, (1–)1.5–3 × (1.5–)3–6 cm, always
wider than long, glabrous, base broadly cordate, margin repand, mucronate at vein endings, apex
rounded; cauline leaves 2–5, simple, petiolate or rarely uppermost subsessile; petiole (0.3–)1–3.5
cm, base not auriculate or rarely with minute auricles to 0.3 × 0.3 mm; leaf blade reniform, 1–2.5
× 1.5–4 cm, base cordate, margin dentate-repand and mucronate at vein endings. Racemes
ebracteate; flowering pedicels 1–2 cm, divaricate, straight, glabrous, stoutish. Sepals ovate or
oblong, 2.5–4 × 1.5–2 mm, glabrous, margin membranous, base of lateral pair saccate; petals
violet, deep magenta, or rarely pale lavender or white, obovate, 7–10 × 3.5–5 mm; median
filament pairs 3.5–5 mm, lateral pair 2–3.5 mm; anthers oblong, 0.9–1.1 mm. Fruits and seeds
not seen.
Flowering: Mar–Apr.
Habitat: sShady wet places, moist stony pastures.
Elevation: 2400−3000 m.
Distribution: China (Guangdong, Sichuan,Yunnan), Myanmar.
Cardamine carnosa Waldstein & Kitaibel, Descr. Icon Pl. Hung. 137, t. 129. 1804; Pterneurum
carnosum (Waldstein & Kitaibel) de Candolle, Syst. Nat. 2: 270. 1821. TYPE: [Croatia,
Velebit Mts.], “inter lapides libros montium clcareorum Croatiam a Dalmatia separantium,”
Kitaibel s.n. (B-WILLD 11974).
Cardamine caroides C. Y. Wu ex W. T. Wang, Acta Bot. Yunnan. 9: 17. 1987. TYPE: China,
Sichuan, Motikonga, Lamashi, 6 Jul 1937, 3800 m, T. T. Yü 6889 (PE!).
Herbs, perennial, glabrous or sparsely to densely pilose with trichomes to 0.7 mm. Rhizomes
thick, to 10 × 5 mm, with numerous stolons. Stems 7–15 cm, erect, simple, thick at base, pilose
or glabrous at base. Rhizomal leaves and lowermost cauline pilose; petiole 1–3 cm; leaf blade
pinnatisect into linear to linear-oblanceolate lobes; terminal lobe obovate in outline, to 8 × 4 mm,
cuneate basally into a petiolule 1–4 mm, deeply trifid or apically three toothed; lateral lobes 1–4
on each side of rachis, 2–8 × 0.5–2 mm, undivided or unequally 2- or 3-fid, margin entire, apex
acute; cauline leaves 2–4, including petiole 1.5–4.5 cm, similar to lowermost cauline leaves but
with fewer and narrower lobes, petiole base not auriculate. Racemes ebracteate, 6–14-flowered;
24
fruiting pedicels ascending or suberect, 0.7–1.5 cm, slender, glabrous, subappressed to rachis.
Sepals ovate or suboblong, 2–2.5 × 0.8–1.5 mm, sparsely pilose, subapically membranous,
lateral pair subsaccate; petals lavender, obovate, 5–6 × 2–2.5 mm, apex obtuse; median filament
pairs 2.5–3.5 mm, flattened and to 0.5 mm wide; lateral pair 1.5–2 mm; anthers oblong, 0.6–0.8
mm; ovules 12–16 per ovary. Fruit (young) ca. 2 cm × 0.8 mm; valves glabrous; style 1–1.5 mm.
Seeds not seen.
Flowering: Jul.
Elevation: about 3800 m.
Distribution: China (Sichuan).
Notes: known thus far only from the holotype.
Cardamine castellana Lihová & Marhold, Taxon 52: 795. 2003. TYPE:
Cardamine changbaiana Al-Shehbaz, Novon 10: 323. 2000. TYPE: China, Jilin, Changbai
Shan, Tianchi, rocky slopes, 2400 m, 1 Aug 1957, Qian Jiaju 580 (holotype, PE!).
Cardamine resedifolia Linnaeus var. morii Nakai, … TYPE:
Herbs, perennial, 2–8 cm, scapose, glabrous throughout. Rhizomes slender, 0.4–0.7 mm in
diam. Stems 2–8 cm, erect, leafless or rarely 1-leaved. Basal leaves rosulate, fleshy; petiole 0.3–
3.5 cm; leaf blade simple or rarely 3-lobed, broadly ovate, subcordate, or oblong, 2–10 × 1.5–8
mm, base cordate or obtuse, margin entire or repand, apex rounded or obtuse; cauline leaves
absent, rarely 1 and petiolate, similar but narrower than basal leaves, petiole base not auriculate.
Racemes ebracteate, 2–5(–7)-flowered; fruiting pedicels erect to erect-ascending, 2–7 mm,
straight. Sepals oblong, 1.3–1.7 × 0.6–0.8 mm; petals white, obovate, 3–3.5 × 1.5–1.8 mm, base
tapering into a claw 0.4–1 mm, apex rounded or subemarginate; filaments 1.4–2 mm; anthers
oblong, 0.5–0.8 mm; ovules 8–12 per ovary. Fruit linear, 1–2 cm × 1.3–2 mm; gynophore 0.3–1
mm; valves smooth, glabrous; style 0.5–2 mm. Seeds brown, oblong to ovate-oblong, 1.2–1.5 ×
0.8–1.1 mm, wingless and not margined.
Flowering: Jul–Aug.
Habitat: rocky slopes.
Elevation: 2400–2500 m.
Distribution: China (Jilin), Korea.
Cardamine chelidonia Linnaeus, Sp. Pl. 2: 655. 1753. TYPE:
Cardamine chenopodiifolia Persoon, Syn. Pl. 2: 195. 1807. TYPE: Uruguay, Montevidio, 1767,
Commerson s.n. (holotype, P-JU; isotype, P!).
Sisymbrium spathulatum Poiret in Lamarck, Encycl. 7: 221. 1806; Arabis spathulata (Poiret) de
Candolle, Syst. Nat. 2: 227. 1821, non Cardamine spathulata Michaux, Fl. Boreal. Amer. 2:
29. 1803. TYPE: Uruguay, Montevidio, 1767, Commerson s.n. (holotype, P-JU; isotype, P!;
fragments, BAA!).
Sisymbrium bellidifolium Poiret in Lamarck, Encycl. 7: 220. 1806; Arabis commersonii de
Candolle, Syst. Nat. 2: 228. 1821, non Cardamine bellidifolia Linnaeus, Sp. Pl. 2: 654. 1753,
nec A. bellidifolia Crantz, Stirp. Austr. Fasc. ed. 1, Fasc. 1, 42. 1762. TYPE: Argentina,
Buenos Aires, Commerson s.n. (holotype, P-JU; isotype, P!).
Heterocarpus fernandezianus Philippi, Bot. Zeit. 14: 641. 1856; Cardamine fernandezianua
Johow, Estud. Fl. Juan Fernand. 110. 1896. TYPE: ……Muñoz Pizzaro (1960) listed Philippi
25
s.n. (SGO-63902) as the type, but that speciemen was collected in September 1872, 16 years
after the description of the species.
Cardamine argentina Spegazzini. TYPE: Argentina, Sierra Ventana, Nov 1895, C. Spegazzini
s.n. (holotype, LP!).
Herbs, annual, sparsely to densely hirsute basally with trichomes to 1 mm, glabescent above,
with chasmogamous and cleistogamous flowers. Stems few to several from base, (0.6−)10−50
cm, erect or decumbent, simple or branched above, hirsute. Basal leaves rosulate, lyrate or
undivided; petiole1−4(−11) cm; leaf blade obovate to spatualte or oblong to oblanceolate,
(1−)2−6(−8) × (0.5−)1−3(−4) cm, sparsely to densely hirsute, sometimes glabrous except for
ciliate margin, base cuneate, margin dentate to crenate, sometimes repand, apex obutse; cauline
leaves petiolate, smaller or same size but gradually reduced in size and becoming subsessile
upward. Racemes ebracteate, 10−30-flowered; fruiting pedicela divaricate to ascending,
4−12(−17) mm. Sepals oblong, 2−2.5 × 1−1.2 mm, caducous; petals white, spatualte to
oblanceolate, 3−5(−6) × 1.5−2 mm, apex obtuse; stamens 6; filaments 2−2.5 mm; anthers oblong
to ovate, 0.4−0.5 mm; ovules (12−)14−20 per ovary. Fruits dehiscent, linear, (1.7−)2−3.5(−4)
cm × 1.5−2.2 mm, thin walled; style 0.5−1(−1.8) mm. Seeds light brown, broadly oblong-ovate,
1.8−2.3 × 1.5−1.8 mm, with a narrow wing 0.1−0.3(−0.4) mm wide. Cleistogamous flowers
solitary, producing geocarpic fruits; sepals to 0.8 mm; petals absent or to 0.5 mm; stamens 2,
median; filaments to 0.6 mm; anthers to 0.2 mm; fruiting pedicels solitary, arising from basal leaf
rosette, (1−)2−5 cm; geocarpic fruits indehiscent, obovate to fusiform, (5−)6.5−10 × 2.4−4 mm,
slightly corky, (1−4)-seeded, without style; seeds oblong, 2.5−3.5 × 2−2.5 mm.
Flowering:.
Elevation: 0−800 m.
Habitat: Damp places.
Distribution: Argentina (Buenos Aires, Chaco, Córdoba, Corrientes, Entre Rios, Formosa,
Misiones), Bolivia (La Paz), Brazil (Rio de Janeiro, Rio Grande do Sul), Chile (Juan
Fernandez Islands), Paraquay, Uruguay (Canelones, Moldonado, Montevideo, San José,
Soriano).
Specimens examined: ARGENTINA. Buenos Aires: Hudson, Parodi 9924 (GH, K, S), Burkart
3952 (SI); Ribera del Rio de la Plata, Punta Larga, Cabrera 2874 (SI); Parana Mini, Burkart
20029 (SI); Dolores, Laguna Sevigné, Correa et 2369 (SI); Balcarce, Cerro de la Virgen, Boelcke
et al. 2402 (BAA); Prtido Balcarce: Sierra de Volcán, Balcarce, Hunziker 3870 (CORD). Partid.
Tigre: Arroyo Gelves, Hunziker 1456.5 (CORD). Chaco. Fontana, Meyer 2322 (LIL). Córdoba:
Punilla, Molinari, Nicora 17637 (SI). Corrientes: Paso de los Libres, laguna Mansa, Goodall &
Tirel 39 (P), Schinini et al. 7200 (SI); Gral. Alvear, rte 40 and rte Aguapey, Lourteig et al. 2836
(P, SI); Mburucuya, Yaguareté-Rincón, Pedersen 790 (K, LP, P, S); Mercedes, 75 km N
Mercedes, Laguna Trin, Ea. Culantrillar, Schinini et al. 11892 (G); Mercedes, Salto Ita Jhase, 9
km SW Boqueron, Teressens et al. 5353 (GH, LIL, MO); Ituzaingó, Isla Apipé Grande, Schinini
& Vanni 15744 (MO); Paso de los Libres, Goodall & Tirel 39 (P); La Cruz, Burkart 7857 (SI).
Dep. Mburucuyá, Estancia Santa Teresa, Burkart 19331 (SI). Dep. Santo Tomé, 7 km E de Gdor.
Virascro, Arroyo Garabí, Schinini & Ahumada 20758 (BACP). Entre Ríos: Gualeguaychi, rio
Paranacito, Burkart et al. 26966 (BACP, K, SI); Concepcion del Uruguay, Troncoso &
Bacigalupo 3209 (BACP, SI), Burkart et al. 28648 (SI). Dep. Concordia, camino de Calabacilla
a Pto. Yeruá, Troncoso et al. 2308 (SI). Formosa: Guaycules, Jörgensen 3423 (LIL, MO, SI).
Misiones: Dep. Iguazu: Parq Nac Iguazú, Hunziker et al. 15481 (CORD), Zardini et al. 854
(LP); Puerto Aguirro, Fangualli 61 (LP). BOLIVIA. Coroico, Bang 2347 (BM, E, F, G, GH, K,
26
MO, NY, W). La Paz: Sud Yungas, Buchtein 178 (BAF, W); Sud Yungas, Puente Villa unos 2
km arriba del río Tamanpaya, Beck 17776 (LPB, MO). BRAZIL. Rio de Janeiro: road between
Alto da Serra and Meio da Serra, 22°33′S, 43°11′W, Smith 1559 (BM, F, GH, K, NY, P, S, UC);
Chemiss du Corcorado, Glaziou 6466 (P). Rio Grande do Sul: Without locality, Saint-Hilaire
1854 (F, P); Porto Aesgre, Linman 339 (G); Parecí, Porto Montenegro, Rambo 42972 (B, LIL);
Vila Manresa, Porto Alegre, Rambo 43476 (B, LIL), Lindman 339 (K, NY, S); Osorio, Rambo
48797 (B, SI); Granja Neugebauer, P. Itapoan, Rambo 48944 (B, LIL), Pelotas, Cost Sacco &
Rambo 824 (F); Morretes, Rio dos Sinos, Rambo 42831 (K, W); San Francisco de Paula
Cambara, Rambo 36544 (S); Sao Leopoldo, Eugenio 163 (NY). CHILE. Juan Fernandez, Reed
89 (SG). PARAGUAY. Doña Juano, Villa-Rica, Balansa 2622 (P). Del Guairá, Iturbe, Montes
12758 (LP). URUGUAY. Doña-Juana, Balansa 2623 (P); Concepcion, Lorentz 1189 (B, G, K,
P, W). Canelones: Santa Lucia, Herter 78937 (F). Moldonado. Puntg Ballena, Hunziker 3989
(BAB). Montevideo: Pocitos/Carrasco, Herter 71215 (G, GH, K, LIL, MO, S, SI, UC); Punta
Yeguas, Osten 22933 (CAS); Carrasco, Legrand 1162 (F); Punta Gorda, Osten 5271 (SI). San
José: Barra Sta. Lucia, Herter 82210 (B). Soriano: Juan Jackson, St. Elena, Gallinal et al. 5254
(MO, NY); Juan Jackson, Monzón-Heber, Gallinal et al. 4388 (MO).
The specific epithets of Sisymbrium bellidifolium and S. spathulatum were published one year
before that of Cardamine chenopodiifolia, but neither of them can be used in Cardamine (see
synonymy above).
Of the approximately 200 species in Cardamine, C. chenopodiifolia is the only species that
produces two fruit types, an indehiscent-geocarpic and dehiscent-aerial. The seeds produced from
subterranean fruits are distinctly heavier and grow faster than those produced from aerial fruits
(Cheplick, G. P. 1983. Differences between plants arising from aerial and subterranean seeds in
the amphicarpic annual Cardamine chenopodiifolia (Cruciferae). Bull. Torrey Bot. Club 110:
442-448.). It has been shown that the subterranean fruiting pedicels produce adventitious roots
(Gamm, R. & F. Weberling. 1986. Sporossbürtige Wurzeln an den unterirdischen Fruchtstielen
bei der amphikarpen Art Cardamine chenopodiifolia Pers. Flora. 178: 19-21). The species also
produces 2-stamened cleistogamous flowers that produce the subterranean fruits and 6-stamened
chasmogamous aerial flowers (Gorczynski, T. 1935. Untersuchungen über Kleistogamie. 4.
Entwicklung der Archesporgeweb und der Befruchtungsvorgang bei Cardamine chenopodiifolia.
Acta Soc. Bot. Pol. 12: 257-274). Fruit and seed anatomy of both areal and subterranean parts
were studied by P. Grimbach (1913. Vergleichende Anatomie verschiedenartiger Frhte und
Samen bei derselben Spezies. Bot. Jahrb. 51(Beibl. 113): 1-52. Cardamine chenopodiifolia: pp.
33-50, figs. 22-31).
Cardamine chenopodiifolia is the only species that produces two fruit types, an indehiscentgeocarpic and dehiscent-aerial. The seeds produced from subterranean fruits are distinctly
heavier and grow faster than those produced from aerial fruits (Cheplick, 1983). The species also
produces 2-stamened cleistogamous flowers that produce the subterranean fruits and 6-stamened
chasmogamous aerial flowers.
Cardamine cheotaiyienii Al-Shehbaz & G. Yang, Harvard Pap. Bot. 3(1): 73. 1998; Hilliella
alatipes (Handel-Mazzetti) Y. H. Zhang & H. W. Li var. macranthus Y. H. Zhang, Acta Bot.
Yunnan. 8: 403. 1986. TYPE: China, Yunnan, Malipo, Guan-gaw, 1000 m, 14 Feb 1940, C.
W. Wang 86836 (holotype, KUN!; isotype, IBSC!).
Herbs, perennial, scapose. Rhizomes ca. 5 mm in diam., with prominent petiolar scars. Stems
(scapes) 15−30 cm, glabrous, leafless. Rhizomal leaves 2, trifoliolate; petiole 5−30 cm,
glabrous; leaflets ovate, 7−18 × 4−5 cm, adaxially glabrous, abaxially sparsely hirsute with thick
trichomes 0.5−0.8 mm, base of terminal leaflet cuneate, that of lateral ones oblique, margin
27
repand to repand-crenate, sparsely ciliate with trichomes 0.1−0.2 mm, apex obtuse to acute;
lateral veins ending with mucros 0.3−0.5 mm; petiolule of terminal leaflet 0.6−1.5 cm, that of
lateral leaflets 2−6 mm; cauline leaves absent. Racemes ebracteate, few flowered, corymbose;
flowering pedicels slender, 2−3.5 cm, glabrous. Sepals ascending, oblong, 5−6 × 2.5−3.5 mm,
erect, slightly saccate at base, glabrous; petals white, oblong, 1.5−1.7 cm × 5−8 mm, erect, apex
rounded; claw 1−2 mm; filaments white, erect, median pairs 6−7 mm, lateral pair 5–5.5 mm;
anthers oblong, ca. 2 mm. Fruits and seeds unknown.
Flowering: Feb.
Elevation: ca. 1000 m.
Distribution: China (Yunnan/Malipo Xian).
Notes: known thus far only from the type collection.
Cardamine chilensis de Candolle, Syt. Nat. 2: 254. 1821. TYPE: Chile, “Dubia tetradynama
siliquosa,” Ruiz & Pavón s.n. (holotype, BM or G). My initial notes from that specimen is
that it probably belongs to C. africana. Therefore, the plant must be checked again.
Cardamine nana Barnéoud in Gay, Fl. Chile. 1: 108. 1846; C. chilensis var. nana (Barnéoud) O.
E. Schulz, Bot. Jahrb. Syst. 32: 445. 1903. TYPE: Chile, Valdivia, C. Gay s.n. (holotype, P!;
isotypes, F!, P!).
Cardamine valdiviana Philippi, Anal. Univ. Chile 27: 314. 1865; C. chilensis var. valdiviana
(Philippi) Reiche, Fl. Chile 1: 94. 1896. TYPE: Chile, Valdivia: San Juan, Oct 1864, F.
Philippi s.n. (lectotype, here designated, SGO-94383!; isotypes, G!, SGO-63908!, in part).
Cardamine solisii Anal. Univ. Chile 27(2): 325. 1865. Chile, Chillán, 1864, Manue Antonio de
Solís s.n. (holotype, SGO-63895!).
Cardamine micropetala Philippi, Anal. Univ. Chile 81: 76. 1892; C. ramosissima Steudel var.
micropetala (Philippi) Reiche, Fl. Chile 1: 97. 1896; C. vulgaris Philippi var. micropetala
(Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 544. 1903. TYPE: Chile, Araucana, R. Philippi
s.n. (lectotype, here designated, SGO-71619!; fragments, BAA!). Muñoz (1960) listed three
type collections, the other two, SGO-49373 and 63906, I have not seen.
Cardamine callitrichoides Spegazzini, Anal. Mus. Nac. Buenos Aires 7: 211. 1902; C.
valdiviana Philippi var. callitrichoides (Spegazinni) O. E. Schulz, Bot. Jahrb. Syst. 32: 446.
1903. TYPE: Argentina, Chubut: Languiñeo, río Carreu-leufú, 1900, N. Illin s.n. (holotype,
LP #12341!; fragemnts, BAA!).
Cardamine chilensis var. angustifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 445. 1903. TYPE:
Chile. Without locality, Apr 1834, Cumming 603 (lectotype, here designated, W!; isotypes,
CGE, K!; fragments, BAA!).
Herbs, perennial or rarely annual, often with thickened, tuberlike stem base, sometimes
rooting from lower nodes, glabrous or with trichomes 0.1−0.2. Stems (2−)4−30(−50) cm, erect,
simple or several from base, simple or branched above, glabrous or sparsely to densely pubescent
near base. Basal leaves and lowermost cauline (0.8−)1.4−4(−5.5) cm, petiolate, not auriculate,
simple or or rarely trifoliolate; petiole (0.4−)0.8−3(−3.5) cm; blade (or terminal leaflet)
(0.2−)0.7−1.5(−2.5) cm × (1−)3−8(−12) mm, oblanceolate to spatulate or obate to ovate, glabrous
or sparsely pubescent, usually minutely ciliate, base cuneate to attenuate or rarely obtuse, margin
entire to repand or rarely crenulate, apex rounded to obtuse; lateral leaflets (when present) ovate
to oblanceolate, considerably smaller than terminal one; middle and uppermost leaves simple,
gradually reduced in size into the racemes. Racemes bracteate throughout or at least along
proximal half, rarely only lowermost few flower bracteat,e many flowered; rachis slightly to
28
strongly flexuous, glabrous; fruiting pedicels (2−)4−10(−17) cm, ascending, slender, straight.
Sepals oblong, 1.2−1.5(−2) × 0.4−0.5 mm, glabrous, caduceus; petals white, oblanceolate,
2.5−3.5(−4) × 0.7−1(−1.5) mm, apex obtuse; filaments 1−2 mm; anthers ovate, 0.1−0.2 mm;
ovules (16−)20−30 per ovary. Fruits 0.7−1.5(−2) cm × 0.7−0.8(−1) mm; style 0.1−0.5(−1) mm.
Seeds light brown, oblong to ovate, 0.7−0.9 × 0.5−0.6 mm, usually with a distal wing ca. 0.1 mm
wide.
Flowering: Aug−Jan.
Elevation: 0−1000 m
Habitat: seepy site, streamside, pond margin, sloughs in dry sandy matorral.
Distribution: Argentina (Chubut, Nuequén, Río Negro, Salta), Chile (Región V, Santiago, VI,
VII, VIII, IX, X).
Specimens examined: ARGENTINA. Chubut: Dep. Futaleufú, Lago Verde, Burkart 19893 (SI).
Dep. Cushamen, valle río Carrileufú, Correa et al. 8956 (BAA, BAB); Parq. Nac. Lago Puelo,
Cusato & Rossow 4040 (BAF). Nuequén: Dep. Huiliches, orilla Lago Paimum, Burkart &
Troncoso 26484 (SI); Lago Paimún, Orilla Norte a 5 km Puesto Gendarmería, Valla et al. 3241
(BAA). Dep. Los Lagos, near Villa la Angostura, Pedersen 1510 (K), Mallo et al. 18175 (BAA);
Lago Nahuel Huapi, Rúgolo 1075 (SI); Bahia Ptol. Manzano, Diem 2962 (BAA), Boelcke 8976
(BAA); Isla Victoria, Boelcke 1796 (BAA); Isla Victoria, Punta Norto, Diem 1892 (LIL, SI); Isla
Victoria, desagüe Laguna Larga, Boelcke 8996 (BAA). Rio Negro: Dep. Bariloche, Park. Nac.
Nahuel Huapi, Isla Victoria, Boelcke 1796 (SI). Dep. Pilcaniyeu, Río Pilcanué, camino Bariloche,
Nicora 3769 (SI). Salta: Dep. La Caldera, camino Jujuy a Salta, Burkart 30460 (SI). CHILE. V:
Catapilco, Philippi 67 (SGO); Las Láunas, [33°06′S, 71°25′W], Jaffuel 3005 (CONC, GH),
Jaffuel 3114 (CONC, GH); Olmué, [32°59′S, 71°09′W], Garaventa 2024 (CONC, SI); Cerro de
La Campana, [32°57′S, 71°08′W], 27 Sep. 1962, Weisser s.n. (CONC); Quilpué, Zöllner 4601
(BACP, SI); Marga Marga, [33°06′S, 71°26′W], Zöllner 7103 (CONC, NA), Jaffuel & Pirion
3056 (GH); Limache, [33°02′S, 71°16′W], Garaventa 2230 (BACP, CONC, SI); Viña del Mar,
[33°02′S, 71°34′W], 30 Aug. 1932, Behn s.n. (CONC); Concón, Pöppig 171 (W); El Pangal,
Garaventa 4090 (BAA, BACP, SI). Santiago: Batuco, [33°12′S, 70°51′W], Garaventa 2206
(BAA, CONC, SI). VI: Caro, 0.6 km W of RR crossing in Alcones on road to Pichilemu,
[34°23′S, 71°43′W], Bliss 547 (CONC); Mine La Leona, Oct. 1935, Grandjot s.n. (MO); Lolol,
[34°44′S, 71°38′W], Barros 2797 (CONC, SI); Colchagua, Lolol, Bliss 754 (CONC); Rancagua,
Bertero 145 (GH, P); Leona, Bertero 146 (G, MO, NY, P, W). VII: Talca, Claude-Joseph 4321
(US); SE Linares along Río Ancoa, along rd to Melado and Medina, 38.2 km upstream from
intersection owoth rd to Peñasco, 35°50-52′S, 71°10-20′W, Taylor & Gereau 10997 (CONC,
MO); Itahue, Fdo. “El Colorado,” [35°08′S, 71°21′W], Garaventa 4539 (BACP, CONC, SI); N
Colbun, 35°38′S, 71°23′W, Bliss 670 (CONC); 10 km E Cauquenes on rd to Parral, 35°59′S,
72°10′W, Bliss 538 (CONC). VIII: Ñuble, 5 km W San Nicholas, 36°29′S, 72°15′W, Bliss 850
(CONC); La Posada, [36°55′S, 73°08′W], Barros 2804 (CONC, SI); 2 km S Escuadron on hwy
160, Lammers, Baeza & Peñailillo 7490 (CONC, MO); Vegas de San Vicente, [36°43′S,
73°07′W], Junge 992 (CONC, SI). IX. Biobio, Cabrero, 5-8 km N Salto El Laja, 37°11′S,
72°23′W, Bliss et al. 2005 (CONC); Malleco, Mininco, [37°47′S, 72°28′W], Montero 5253
(CONC, SI); 5-6 km SE Minico on rd to Collipulli, Bliss 864 (CONC); Araucania, Nov. 1887,
Philippi s.n. (SGO); E Pucón, from the Río Turbio along rd to Termas de Huife, 39°20′S,
71°45′W, Taylor & Taylor 10870 (MO). X: Valdivia, Hohenacker 275 (G, P), Buchtein s.n.
29
(US); Talcahuano, Pöppig 168 (P, W); Quinchilca, [39°52′S, 72°46′W], Hollermayer 870
(CONC, SI). Bío Bío, Antuco, Barros 2803 (SI); Llanquihue, Peulla, Pennell 12669 (GH); San
Miguel, Barros 798 (BACP).
Cardamine chilensis is highly variable in habit, and under favorable conditions, the plants
perenniate, and the stem base becomes only slightly to considerably indurated to tuberlike and
often several stems are produced. This very aspect of the stem base was used by Schulz (1903)
and Boelcke and Romanczuk (1984) as the basis for recognizing C. valdiviana as separate from
the slender-based C. chilensis. In fact, the type of C. valdiviana is a single-stemmed plant only
very slightly thickened at the stem base. No other differences can be found in the numerous
collections examined in this complex.
There is considerable variation in the racemes, and the variation ranges from fully bracteate
racemes to those basally bracteate or rarely only the lowermost few flower bracteate. Cardamine
solisii is indistinguishable from C. chilensis in every aspect except for the nearly ebracteate
raceme. Indeed, only the lowermost flower of the type of C. solisii is bracteate, but there are
collections of C. chilensis with two, three, or more bracts. Should these be considered as distinct
species? My answer is no. Because of the lack of any diffences among C. chilensis, C.
valdiviana, and C. solisii in leaf, flower, and fruit morphology, I am recognizing a single species
undividable into infraspecific taxa, as Schulz (1903) and Boelcke and Romanczuk (1984) did.
Cardamine circaeoides J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5: 144. 1861. TYPE:
India, Sikkim, 5000–7000 ft, J. D. Hooker s.n. (holotype, K!; isotypes, B!, P!).
Cardamine agyokumontana Hayata, Ic. Pl. Formosa. 3: 19. 1913. TYPE: Taiwan (Formosa), ….
(holotype, …; photos, A, US).
Cardamine circaeoides var. diversifolia O. E. Schulz,
Cardamine heterandra J. Z. Sun & K. L. Chang, Tawania 41: 113−116. 1996. TYPE. China,
Gansu, Wudu County, Pandi Xiang, 1,900 m, 19 Apr 1993, Baishuijiang Team 2209
(holotype, LZU).
Cardamine insignis O. E. Schulz, Bot. Jahrb. Syst. 32: 439. 1903. TYPE: China, Yunnan, 6000
ft., A. Henry 13,090 (holotype, B!; isotypes, E!, K!, MO!).
Cardamine macrocephala Z. M. Tan & S. C. Zhou, J. Sichuan Univ. (Natural Sci.) 33(5): 601.
1996. TYPE: China, Sichuan, Tianquan County, Erlong Mountain, Xingou, 6 Jun 1959,
Tianquan Expedition 665 (holotype, CDBI).
Cardamine reniformis Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25(19): 50. 1908.
Cardamine violifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 440. 1903. TYPE: China, Hupeh, near
Ichang, A. Henry 3298 (holotype, B!; isotypes, E!, GH!, K!, P!, US!).
Cardamine violifolia var. diversifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 440. 1903. TYPE:
China, Hupeh, near Ichang, A. Henry 3298 (same as above)(holotype, B!; isotype, GH!).
Cardamine violifolia var. pilosa K. L. Chang & H. L. Huang, Taiwania 41: 115. 1996. TYPE:
China, Gansu, Wen Xian, Tielou Xiang, 2,200 m, 19 Sep 1993, Baishuijiang Team 1572
(holotype, LZU).
Herbs, perennial, (5–)9–36(–45) cm, sparsely to densely pilose at least on leaves. Rhizomes
slender, sometimes with a few stolons. Stems slender, erect or decumbent, simple or rarely
branched above middle. Basal leaves not fleshy, simple or rarely 2–4-foliolate; petiole 1–9(–12)
cm, glabrous or rarely ciliate; leaf blade or terminal leaflets cordate to ovate, rarely suborbicular,
(0.7–)1.5–5.5(–6.7) × (0.7–)1.5–4.3(–5) cm, always longer than wide, pilose or glaborous, base
cordate, sometimes obtuse, subtruncate, or subcuneate, margin often repand-crenate, sometimes
coarsely crenate, crenulate, or subentire, often distinctly mucronate at vein endings, apex obtuse;
lateral leaflets (when present) petiolulate or sessile, much smaller than terminal one; cauline
30
leaves 1–4, simple or rarely one 2- or 3-foliolate, petiolate or rarely uppermost subsessile; petiole
(0.4–)1–5(–6) cm, with basal auricles 0.4–2 × 0.2–0.4 mm; leaf blade similar to that of basal
leaves, cordate, sometimes ovate to ovate-lanceolate, 1–3.5(–6) × 1–3(–3.5) cm. Raceme
ebracteate, many flowered; fruiting pedicels 3–12(–15) mm, ascending, divaricate, or reflexed,
sometimes secund, glabrous, slender. Sepals ovate or oblong, 2–3.5 × 0.8–1.5 mm, glabrous,
margin often membranous, base not saccate; petals white, spatulate, (4–)5–7(–8) × (1.5–)2–2.5(–
4) mm; median filament pairs (2.5–)3.5–5 mm, lateral pair (2–)2.5–3.5 mm; anthers oblong, 0.5–
1 mm; ovules 20–42 per ovary. Fruit linear, 1.3–3(–3.3) cm × 0.8–1.2(–1.5) mm; valves
torulose, glabrous; style (0.5–)1–2 mm. Seeds brown, ovate to broadly oblong, 0.8–1.1 × 0.6–0.9
mm; wingless.
Flowering: Feb–Jul.
Habitat: ravines, along streams and ditches, rocky places, mixed woods, moist pastures,
roadsides, forests.
Elevation: 400–3300 m.
Distribution: China (Gansu, Guangdong, Guangxi, Hunan, Sichuan, Taiwan, Yunnan), India
(Sikkim), Laos, Myanmar, Thailand, Vietnam.
Cardamine clematitis Shuttleworth ex A. Gray, Proc. Amer. Acad. Arts 15: 45. 1880. TYPE:
Herbs, perennial, glabrous throughout. Rhizomes slender, cylindrical, 1–3 mm in diam.;
stolons absent. Stems (0.8–)1–2.5(–3.5) dm, erect, simple or rarely few branched above,
glabrous. Rhizomal leaves simple or 3-foliolate, (1.5–)3–8 cm; petiole (1–)2–6 cm; blade or
terminal leaflet reniform to cordate, (0.5–)1–2 cm, petiolule to 1 cm, base cordate, margin entire
or 3-lobed, not puberulent; lateral leaflets much smaller than terminal one, subsessile, margin
same as terminal leaflet; cauline leaves 3–7, 3-foliotate or uppermost simple; petiole 0.7–3.5 cm,
base auriculate, the auricle 0.7–5 mm; terminal lobe broadly ovate to suborbicular or reniforem,
1.5–4 × 1.5–3.5 cm, often 3–5-lobed, the lobe apex minutely apiculate, glabrous along margin,
petiolule 0.5–1 cm; lateral leaflets oblong to ovate or oblong, sessile or on petiolule to 0.5 cm.
Racemes ebracteate; fruiting pedicels divaricate-ascending, (0.7–)1–1.7 cm. Sepals oblong, 2.5–
3 × 1–1.5 mm, erect, base of lateral pair not saccate; petals white, oblanceolate, 6–8 × 2–3 mm,
not clawed, apex obtuse to subemarginate; median filament pairs 3.5–4 mm, lateral pair 2–2.5
mm; anthers ovate, ca. 0.7 mm. Fruit linear, (1.5–)2–3.5–(4) cm × 1.3–1.7 mm; valves glabrous;
style 2–4 mm; ovules and seeds 10–16 per fruit. Seeds brown, oblong, 1.7–2 × 1–1.2 mm.
Flowering: May–Jun.
Habitat: wet areas, springs, moist slopes.
Elevation: 1300–1800 m.
Distribution: United States (Alabama, Florida, Georgia, North Carolina, Tennessee, SW
Virginia).
Cardamine concatenata (Michaux) O. Schwarz, Feddes Repert. Sp. Nov. Regni Veg. 46: 188.
1939; Dentaria concatenata Michaux, Fl. Bor.-Amer. 2: 30. 1803. TYPE:
Cardamine laciniata var. integra O. E. Schulz, Bot. Jahrb. Syst. 32: 349. 1903; Dentaria
laciniata var. integra (O. E. Schulz) Fernald, Rhodora 10: 84. 1908. TYPE:
Cardamine laciniata var. lasiocarpa O. E. Schulz, Bot. Jahrb. Syst. 32: 349. 1903. TYPE:
Dentaria laciniata Muhlenberg ex Willdenow, xxx; Cardamine laciniata (Muhlenberg ex
Willdenow) A. Wood, Amer. Bot. & Fl. 38. 1870, not C. laciniata Steudel, Nomencl. ed. 2,
pt. 1: 281. 1840. TYPE:
Dentaria laciniata var. alterna Farwell, Amer. Midl. Naturalist 12: 58. 1930. TYPE:
31
Dentaria laciniata var. coalescens Fernald, Rhodora 40: 421. 1938. TYPE:
Dentaria laciniata var. latifolia Farwell, Amer. Midl. Naturalist 12: 58. 1930. TYPE:
Dentaria laciniata var. opposita Farwell, Amer. Midl. Naturalist 12: 58. 1930. TYPE:
Herbs, perennial, sparsely pubescent or sometimes glabrous. Rhizomes fleshy, moniliform,
segments fusiform, 2–10 mm in diam.; stolons absent. Stems (1–)2–4(–5.5) dm, erect, simple,
glabrous or pubescent distally. Rhizomal leaves 3-foliolate, (7–)10–20(–30) cm; petiole (4–)7–
18(–25) cm; terminal leaflet oblong, lanceolate, oblanceolate, or linear, 2.5–6 cm, sessile, base
cuneate, margin coarsely dentate to incised or laciniate, sometimes 3-lobed and the lobes toothed
to incised, rarely entire, puberulent with trichomes 0.2–0.3 mm, or not; lateral leaflets about as
large as or smaller than terminal leaflet, sessile, margin same as terminal leaflet; cauline leaves (2
or) 3, 3-foliolate, whorled to opposite or rarely alternate, similar in morphology to rhizomal
leaves; petiole 1–)1.5–6(–7.5) cm, base not auriculate; terminal leaflet lanceolate, linear, or
oblanceolate, (3–)4–10(–12) × (0.3–)0.5–2(–2.5) cm, coarsely dentated to incised or rarely
subentire, minutely puberulent along margin, subsessile or with petiolule to 3 cm; lateral leaflets
similar to terminal one, sometimes smaller, sessile. Racemes ebracteate; fruiting pedicels
ascending to divaricate, (0.6–)1–2.7(–3.3) cm. Sepals oblong, (4–)5–8 × 2–4 mm, erect, base of
lateral pair slightly saccate; petals white to pale pink, oblanceolate, (8–)10–20 × (3–)4–7(–9)
mm, short clawed, apex rounded; median filament pairs 8–12 mm, lateral pair 6–8 mm; anthers
oblong-linear, 1.5–2.5 mm. Fruit linear-lanceolate, (2–)2.5–3.8(–4.8) cm × 1.5–3 mm; valves
glabrous or sparsely pubescent; style (2–)5–9(–12) mm; ovules and seeds 10–14 per fruit. Seeds
brown, oblong, 1.6–3 × 1.8–2 mm. 2n = 128–256.
Flowering Feb–May.
Habitat: wooded bottoms and bluffs, rich woods, limestone cliffs and outcrops, rocky banks,
mesic forests, moist areas with leaf litter, floodplain woods.
Elevation: 0–1000 m.
Distribution: Canada (Ontario, Quebec), United States (Alabama, Arkansas, Connecticut,
Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky,
Louisiana, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, E
Nebraska, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oklahoma,
Pennsylvania, South Carolina,Tennessee, E. Texas, Vermont, Virginia, West Virginia,
Wisconsin).
Cardamine conferta
Cardamine constancei Detling, Madroño 3: 176. 1935. TYPE:
Herbs, perennial, glabrous or sparsely hirsute. Rhizomes slender, cylindrical, to 2 mm in
diam.; stolons absent. Stems 1.5–5 dm, erect, simple, glabrous or sparsely hirsute near base.
Rhizomal leaves absent; cauline leaves 4–7, crowded above middle of stem, simple, (5–)7–15(–
20) cm; petiole (0.5–)1–4(–5) cm, base not auriculate; leaf blade broadly ovate or ovatelanceolate to ovate-elliptic, rarely obovate, 5–13 × 2–6.5 cm, minutely pubescent along margin,
base cuneate, margin coarsely serrate with small apiculae, or repand to undulate, apex acute.
Racemes ebracteate; fruiting pedicels divaricate-ascending to suberect, 1–2.2 cm. Sepals oblong,
6–8 × 2–2.5 mm, erect, base of lateral pair slightly saccate; petals pink, oblanceolate, 15–28 × 5–
8 mm, claws to 10 mm, apex rounded; median filament pairs 8–10 mm, lateral pair 4–6 mm;
anthers linear, 2–3 mm. Fruit linear, 2.5–3.5(–5) cm × 1.9–2.1 mm; valves glabrous; style 2–3.5
mm; ovules and seeds 12–16 per fruit. Seeds brown, oblong, 2–2.5 × 1.2–1.5 mm.
Flowering: May–Jun.
32
Habitat: moist cliffs, wooded creek bottoms, shaded draws, hillsides, moist woods, mixed
conifereous forests, granitic soils.
Elevation: 400–600 m.
Distribution: United States (N Idaho/Cleawater, Idaho, Kootenai, Nez Perce, and Shoshone
counties).
Cardamine cordata Barnéoud in Gay, Fl. Chile. 1: 109. 1846. TYPE: Chile, Coquimbo, 1839,
C. Gay s.n. (holotype, P!; isotypes, F!, K!, P!; fragments, BAA!).
Cardamine decumbens Barnéoud in Gay, Fl. Chile 1: 109. 1846; C. cordata Barnéoud var.
decumbens (Barnéoud) O. Schulz, Bot. Jahrb. Syst. 32: 430. 1903. TYPE: Chile. Coquimbo:
Cordillera de Guanta, 2490 m, 1838, C. Gay 329 (holotype, P!; possible isotype, P!;
fragments, BAA!)
Cardamine calbucana Philippi, Anal. Univ. Chile 41: 668. 1872; C. cordata prol. calbucana
(Philippi) O.E. Schulz, Bot. Jahrb. Syst. 32: 429. 1903. TYPE: Chile. Volcán Calbuco, Jan.
1872, Carlos Juliet s.n. (holotype, SGO-49393!; fragment, BAA!). Muñoz (1960) also listed
SGO-63898 as the type collection, but I have not seen that.
Cardamine andina Philippi, Anal. Univ. Chile 81: 71. 1892; Cardamine nivalis Gillies var.
andina (Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 519. 1903. TYPE: Chile. Cordillera de
Las Condes, 7-8000 ft, Jan. 1891, Edwyn Reed s.n. (lectotype, here designated, SGO-63884!).
Muñoz (1960) listed two other sheets, SGO-63912! and SGO-71625!, as part of the type
collection. The one I designated as the lectotype agrees best with the original description.
Cardamine monticosa Philippi, Anales Univ. Chile 81: 72. 1892. TYPE: Chile. Prov. Colchagua,
Las Vinas, Jan 1872, Philippi s.n. (lectotype, here designated, SGO-49335!). Muñoz (1960)
listed three sheets as part of the type collection of this species, of which SGO-49335! is C.
tenuirostris and does not correspond morphologically with the original desription of C.
monticosa. I have not seen the third sheet, SGO-49322 (fragments, BAA!).
Cardamine peteroana Philippi, Anal. Univ. Chile 81: 74. 1892. TYPE: Chile. Curicó: Thermas
de Peteroa, Manuel Vidal s.n. (holotype, SGO-…). not seen, and no sheets listed by Muñoz
(1960).
Cardamine rostrata Grisebach var. alpina Chodat & Wilczek, Bull. Herb. Boiss. Ser. 2, 2: 288.
1902. TYPE: Argentina. Cajon del Burro, 3000 m, Wilczek 431 (holotype, G, not seen;
fragments, BAA!).
Perennial herbs, glabrous throughout. Rhizomes not tuberous, short, slender, not scaly,
rarely swollen at the well-spaced nodes. Stems single from rhizome, (3−)5−33(−55) cm tall,
erect or subdecumbent, glabrous, simple or rarely branched above, with 2−8 croweded or
rarely well-spaced leaves. Basal leaves not rosulate, they and lowermost cauline leaves simple
or rarely with 1 or 2 pairs of lateral leaflets; petiole 1−4(−8) cm long; leaf blade or terminal
leaflet orbicular to cordate or reniform, (0.5−)1−3.5(−4) × (0.8−)1.5−4(−4.5) cm, often wider
than long, palmately veined, always fleshy, base often deeply cordate, repand to crenate or
obtusely angled, rarely veins ending in a minute mucro, petiolule (if leaves compound)
0.5−1.5 cm long; lateral leaflets (when present) sessile or less frequently on petiolules 0.5−3
mm long, cordate or oblong to ovate, 0.5−12(−25) × 3−7(−17) mm, base usually oblique;
middle and upper cauline leaves simple or rarely, 3−7-foliolate leaflets, smaller and narrower
than lowermost leaves, margin entire, repand or obtusely dentate. Racemes corymbose,
ebracteate, elongated slightly or moderately in fruit; rachis straight; fruiting pedicels
(0.2−)0.5−1(−1.8) cm long, ascending erect, forming a straight line with fruit, appressed to
rachis, stout, glabrous. Sepals oblong, 3−4(−4.5) × 1.5−1.8 mm, erect, caducous, glabrous.
33
Petals white, oblanceolate to obovate, 7−9 × 2.5−4 mm, apex obtuse. Filaments 3.5−5 mm;
anthers oblong, 0.7−1 mm. Ovules 20−38 per ovary. Fruits linear, (1.8−)2−3.5 cm × 1.2−1.5
mm, slightly attenuate to apex; style 0.7−2(−3) mm long. Seeds brown, oblong to broadly
ovate, 1−1.5 × 0.8−1 mm, sometimes narrowly winged at apex.
Fl.: Oct−Feb.
Alt.: 700−2900 m
Habitat: Brooks, springs, wet areas, meadows, moist slopes along rivlets, wet flushes near
springs, stunted Nothofagus forest to base of snow.
Distribution: Argentina and Chile.
Specimens examined: ARGENTINA. Cajon del Burro, Gerk 109 (SI). Chubut: Dep.
Languiñeo, Rio Tecke, Ea. Pampa Chica, Soriano 3817 (BAA); Co. Colorado, Corcovado,
Krapovickas 4083 (BAA, BAB). Dep. Rio Senguerr, Lago Blanco, Koslowsky 12492 (NA);
Laguna Blanca, 45°52′S, 71°15′W, Koslowsky 45 (SI). Dep. Tehuelches, Lago Verde, Soriano
4255 (BAA). Mendoza: arroyo de loas Alverjalitos, Kurtz 5822 (CORD, LP). Rio Salado
Valley, 335°S, Böcher et al. 1019 (BAA, MO). Dep. Malargüe, Ruta Prov. 224, camino a Po.
Pehuenche, Puesto Rojas, 35°57′S, 70°21′W, Prina et al. 2047 (SI, SRFA); Arroyo Negro, Kurtz
5737 (CORD, LP). Dep. San Rafael, Arroyo Grande frente de Los Molles, Sleumer 538 (LIL);
Portezuelo Ancho an Camino a Valle Hermoso, Leal 24582 (BAA). Neuquén: Dep. Chos Malal,
Arroyo Dominyo, 36°44′S, 70°23′W, Boelcke et al. 11304 (BAA, BAB, SI); Los Tábonos,
Boelcke et al. 11339 (BAA, BAB, SI). Dep. Minas, Lagunas Epu-Lauquén, Boelcke et al. 10902
(MO, SI); Vegas del Pelán, Boelcke et al. 11131 (BAA, BAB, LIL, MO, SI); Piedra del Gallo,
36°42.5′S, 70°30′W, Boelcke et al. 11396 (BAA, SI); Sierra de Cochicó, cajón de la Botica,
36°21-22′S, 70°34-36′W, Boecke et al. 14113 (BAA, BACP, SI); cajón del Portillo, 36°12′S,
70°36′W, Boecke et al. 14161, 14185 (BAA, BAB, BACP, SI); confluence rós Pichi-Neuquén
and Neuquén, cerro de las Yeguas, 36°35′S, 70°46′W, Boelcke et al. 13778 (BAA, BAB, BACP,
SI); Colunco, Ammann 50 (F); Lapala, Mallin, Ammann 50b (F); Cord. Del Viento, Arroyo
Mamanque, Patore 31 (SI); Parq. Nac. Nahuel Huapi, Refugio Co. Colorado, E de Punto,
Boelcke 9028 (SI). Dep. Minasa, cruzada de Tricao Malal al Cajón de Butaló, 36°58′S, 70°30′W,
Boelcke et al. 11564 (BAA, BAB). Dep. Los Lagos, Parq. Nac. Nah. Huapi, refugio Co.
Colorado, Boelcke 9025 (BAA); Ea. Fortín Chacabuco, Correa et al. 3785 (BAA, BAB). Dep.
Ñorquín, Copahue, Rossow et al. 859 (BAB, BACP). Dep. Picunches, Pino Hachado, Rossow
1140 (BAB, BACP). Río Negro: Cerro López, 41°, Böcher et al. 1745 (BAA, MO), Boelcke
1838 (SI); Bariloche, Cerro López, Rassow & Gómez 370 (BACP); Cerro Ñireco, Mallui, 6 Jan.
1947, Beruasconi s.n. (SI); Parque Nac Nahuel Huapi, Cerro Tronador, Boelcke & Correa 5742
(BAB, SI); Parq. Nac. Nahuel Huapi, Pampa Linda, Ventisqueros del Tronador, Boelcke et al.
10454 (BAA, SI). Dep. Pilcaniyeu, Los Juncos, Ea. San Ramón, Port. Lana Grande, Boelcke &
Correa 11737 (BAA); Valle de Comallo, Vallerini 2794 (SI); Co. Tronador, garganta del Diablo,
16 Jan 1988, Valla et al. sn. (BAA); Cerro Lopez, Parodi 11491 (BAA). Santa Cruz: Lago
Buenos Aires, near Perito Moreno, Ager 436 (US). CHILE: IV: Dep. Ovalle, Quebrada del
Toro, Pabellon, 20 km E Hurtado, Hacienda El Bosque, Wagenknecht 18491 (F, G, GH, MO,
UC), Wagenknecht 4091 (CONC, SI). V: Los Andes, de Portillo rd, base de Caracoles, Charpin
& Lazare 23820 (G). Santiago: Paso de Jorquera, Farellones, [33°21′S, 70°19′W], Garaventa
4517 (CONC, SI); Los Perales, [33°45′S, 71°06′W], Montt 233 (CONC); near Rio Colorado,
Hastings 430 (US); Piuquencillos, Valle del Rio Colorado, [33°35′S, 70°13′W], Pisano et al.
1662 (CONC). VI: Cord. Colchagua, 1870, Philippi s.n. (G); Adnes of Rancagua, [33°54′S,
34
70°28′W], Montt 288 (CONC). VII: Thermas de Longavi, Jan. 1888, Schoenemann s.n. (SGO);
Cordl. Talca, Feb. 1879, F. Philippi 68e (SGO); Cord. del Peuco, [33°56′S, 70°30′W], Cádiz s.n.
(SGO); Peteroa, [35°15′S, 70°34′W], Grandjot 602 (CONC, SI), Philippi s.n. (SGO); Laguna de
Teno, [35°10′S, 70°35′W], Marticorena & Matthei 908 (CONC), Marticorena et al. 28 (CONC);
Laguna del Maule, [36°00′S, 70°30′W], Schlegel 3540 (CONC). VIII: Laguna del Loja,
[37°21′S, 71°19′W], Garaventa 4626 (BACP), Montero 6127 (CONC); Baños de Chillan,
Werdermann 1313 (CAS, CONC, E, F, G, GH, LIL, MO, NY, SI, UC, US), Pennell 12441 (F,
GH, NY, US); Cordillera de Chillán, 1855, Germain (G, SGO); Santa Barbara, 1839, Gay s.n.
(SGO); Termas de Chillán, [36°54′S, 71°31′W], Pfister 8754 (CONC), Jaffuel 2005 (CONC);
Valle de las Nieblas, Feb. 1892, Philippi s.n (SGO); Cordillera de Los Andes, rd to Shangrila,
36°52′S, 71°27′W, Gardner & Knees 6827 (E, MO); Las Fumarolas and Valle de las Nieblas,
36°54′S, 71°23′W, Tayloer et al. 10297 (CONC). IX: Volcán Llaima, [38°43′S, 71°43′W],
Gunckel 12281, 13905 (CONC), Montero 6834 (CONC); Termas Rio Blanco, Montero 3397
(GH); Termas de Alpehue, [38°54′S, 71°37′W], Montero 10846 (CONC); Curacautín, Cord. de
los Andes, Parque Nacional Conguillío, 38°36′S, 71°36′W, Brownless et al. 990 (MO); Malleco,
Sierra Nevada, Laguna Conguillío, [38°36′S, 71°35′W], Schlegel 7152 (CONC). X: Volcán
Choshuenco, [39°56′S, 72°04′W], Hollermayer 799 (CONC); Dep. t. Puerto Varas, Peulla,
[41°06′S, 72°02′W], Zollitsch 269 (CONC); Parque Nacional Puyehue, Antillanca, 40°46′S,
72°12′W, Gardner & Knees 3998 (CONC, E), Schlegel 6770, 7301 (CONC), Sparre &
Constance 10780 (CONC); Paso Payehue, [40°40′S, 71°55′W], Sparre & Constance 10833
(CONC); Palena, Las Escalas, Futaleufú, 43°12′S, 71°50′W, Hildebrand-Vogel 32 (CONC). XI:
Estanci Coyhaique Alto, near Cerro Coyhaique, Santesson 1287 (S). Prov.??: Valle de las agues
Calientes, 27 Feb. 1862, Philippi s.n. (SGO); Cajón de Azufre, Feb. 1891, Albert s.n. (SGO);
Cajón del Calro, Feb. 1891, Albert s.n. (SGO); Al pié de la Quebrada Ventana, Garaventa 3131
(BACP, CONC, SI).
A comparison of the types of Cardamine calbucana and C. monticosa reveals that they are
indistinguishable from that of C. cordata. Those of C. andina and C. decumbens represent robust
forms of the species that are erect or decumbent, respectively. The C. andina type has sessile and
somewhat decurrent leaflets of upper leaves, but this aspect, though rare in typical C. cordata,
does not merit the creation of an inDep. endent taxon. As for C. decumbens, it might deserve to
be recognized as an ifraspecific taxon of C. cordata, as was done by Schulz (1903), but in the
absence of fieldwork, it would be better to retain it as a synonym.
Cardamine cordifolia A. Gray, Mem. Amer. Acad. Nat. Sci. (Pl. Fendl.) 4: 8. 1849. TYPE:
United States, New Mexico, Santa Fe Co., margin of Santa Fe Creek, 15 May–16 Jun 1847,
A. Fendler 28 (holotype, GH!; isotypes, K!, MO!, NY!, PH, UC!).
Cardamine cardiophylla Rydberg, Bull. Torrey Bot. Club 28: 280. 1901, not Greene, Fl.
Francisc. 266. 1891; C. cordifolia var. cardiophylla O. E. Schulz, Bot. Jahrb. Syst. 42: 438.
1903; C. infausta Greene, Pittonia 4: 307. 1901. TYPE: United States, Colorado, Lake Co.,
Tennessee Pass, 27 Jun 1900, G. E. Osterhout 2178 (holotype, NY!; isotypes, GH!, RM).
Cardamine cordifolia var. diversifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 438. 1903. TYPE:
United States, New Mexico, Santa Fe Co., Santa Fe Canyon, 9 miles E of Santa Fe, 8000 ft, 2
Jun 1897, A. A. & E. G. Heller 3647 (holotype, B; isotypes, DS!, MO!, NY!).
Cardamine cordifolia var. incana A. Gray ex M. E. Jones, Proc. Calif. Acad. Sci. II, 5: 620.
1895; C. incana (A. Gray ex M. E. Jones) A. Nelson, Bot. Gaz. 42: 50. 1906. TYPE: United
35
States, Utah, Piute Co., Marysvale, 7000 ft, 1 Jun 1894, M. E. Jones 5341a (holotype, POM;
isotype, MO!).
Cardamine cordifolia var. pubescens A. Gray ex O. E. Schulz, Bot. Jahrb. Syst. 32: 439. 1903.
TYPE: United States, Utah, Wayne Co., S slope of Thousand Lake Mt., 8000 ft, 14 Jul 1875,
L. F. Ward 396 (lectotype designated by Welsh (1982: 162), US!; isolectotype, GH!, NY!).
Cardamine lyallii S. Watson, Proc. Amer. Acad. Arts 22: 466. 1887; C. cordifolia subsp. lyallii
(S. Watson) O. E. Schulz, Bot. Jahrb. Syst. 32: 438. 1903; C. cordifolia var. lyallii (S.
Watson) A. Nelson & J. F. Macbride, Bot. Gaz. 61: 31. 1916. TYPE: (lectotype should be
designated).
Cardamine uintahensis F. J. Hermann, Rhodora 36: 410. 1934. TYPE: United States, Utah,
Summit Co., muddy bank of Mill Creek, SW base of Mt. Elizabeth, 6 miles SE of Goodman
Ranch, Bear River Valley, 8500 ft, F. J. Hermann 5894 (holotype, GH!).
Herbs, perennial, glabrous or densely puberulent. Rhizomes slender or stout, cylindrical, 1.5–
3 mm in diam.; stolons absent. Stems 2–7(–10.2) dm, erect, simple or branched above, glabrous
or sparsely to densely puberulent near base or throughout. Rhizomal leaves absent or rarely
present, simple, 5–15 cm; petiole 2.5–12 cm; leaf blade reniform to cordate, 1.5–4.5(–6) cm;
cauline leaves (3–)5–17(–23), simple or very rarely lowermost 3-foliolate; petiole 1–6.5(–8.5)
cm, base not auriculate; leaf blade somewhat fleshy, reniform, deltoid-cordate, or ovate-cordate,
(1–)2–7.2(–9.7) × 1–5.5(–8.5) cm, glabrous or sparsely to densely pubescent, base cordate or
truncate, margin crenate to slightly sinuate, veins ending in small apicula, puberulent or not.
Racemes ebracteate; fruiting pedicels divaricate to ascending, (0.7–)1–2 cm. Sepals oblong, 2.5–
4.5 × 1.5–2 mm, erect, base of lateral pair slightly saccate; petals white, broadly obovate, 7–12 ×
4–6 mm, with a claw to 6 mm, apex rounded to subtruncate or emarginate; median filament pairs
3.5–5 mm, lateral pair 2–3.5 mm; anthers oblong,1–1.5 mm. Fruit linear, (2–)2.5–3.7(–4) cm ×
1.2–2 mm; valves glabrous or sparsely puberulent; style 0.5–3(–6) mm; ovules and seeds 14–24
per fruit. Seeds brown, oblong, 1.6–2 × 1–1.3 mm. 2n = 24.
Flowering: May–Aug.
Habitat: streambanks, springs, shady gullies, creek bottoms, lake shores, ponds, cold springs,
meadows, moist hillsides, mossy areas, alpine streams, mixed coniferous forests.
Elevation: 600–3600 m.
Distribution: Canada (British Columbia), United States (Arizona, California, Colorado, Idaho,
New Mexico, Nevada, Oregon, Utah, Washington, Wyoming).
Cardamine corymbosa J. D. Hooker, Fl. Antarct. 1: 6. 1844; C. hirsuta Linnaeus var.
corymbosa (J. D. Hooker) J. D. Hooker, Handb. New Zeal. Fl. 12. 1864. TYPE:
Cardamine crassifolia Pourret
Cardamine debilis Banks ex de Candolle, Syst. Nat. 2 : 265. 1821; not C. debilis D. Don, Prodr.
Fl. Neal. 201. 1825; Cardamine hirsuta Linnaeus var. debilis (Banks ex de Candolle) J. D.
Hooker, Handbook New Zeal. Fl. 12. 1864. TYPE: New Zealand,.. (holotype, xx).
Sisymbrium heterophyllum G. Forster, Prodr. 46. 1786; Cardamine heterophylla (G. Forster) O.
E. Schulz, Bot. Jahrb. Syst. 32: 487. 1903, not C. heterophylla (Nuttall) A. Wood Amer. Bot.
& Fl. 38. 1870; not Host, Syn. P. Austral. 366. 1797; not Lapeyrouse, Hist. Abr. Pl. Pyr. 377.
1813; not Bory, Ann. Sci. Gen. Phys. 3: 6. 1820; not W. J. Hooker, Comp. Bot. Mag. 1: 273.
1835. TYPE: New Zealand,.....(holotype, ).
36
Cardamine delavayi Franchet, Bull. Soc. Bot. France 33: 397. 1886. TYPE: China. Yunnan:
near Mo-so-yun, Lankong, 2 Apr 1885, Delavay 1838 (holotype, P!; isotypes, P!, US!).
Herbs, perennial, 15–45, sparsely pilose on leaves and at nodes, or glabrous throughout.
Rhizomes slender and much elongated, with a few stolons and no bulbils. Stems simple at base,
rarely branched above, erect, strongly flexuous; nodes well spaced, lowermost internodes to 9
cm. Rhizomal leaves simple; petiole 3–12 cm; leaf blade reniform, cordate, or suborbicular, 1.2–
2.5 × 1.5–3 cm, palmately veined, base cordate, margin entire or obscurely 5-lobed, endings of
principal veins and leaf apex mucronate; cauline leaves 2–5, trifoliolate; petiole of lowermost
and middle ones 2–5 cm, not auriculate and sometimes ciliate at base; terminal leaflet broadly
oblong or narrowly obovate, 0.7–2 × 0.7–1.5 cm, with a petiolule to 4 mm, margin entire, apex
mucronate; lateral leaflets similar to terminal, slightly smaller. Uppermost cauline leaves
trifoliolate; petiole 1–3.5 cm, not auriculate at base; terminal leaflet narrowly oblong or linear,
sessile, 1–3 cm × (1–)2–4(–5) mm; lateral leaflets smaller. Racemes ebracteate, 7–15-flowered;
fruiting pedicels ascending, (1–)1.2–2(–2.3) cm, straight. Sepals ovate, 2–3 × 1–1.5 mm, margin
membranous; petals white, broadly obovate, 5–7 × 2.5–4 mm, apex rounded; median filament
pairs 2.5–4, not flattened; lateral pair 1.5–3 mm; anthers linear-oblong, 1–1.3 mm; ovules 10–20
per ovary. Fruit linear, 2–3 cm × 1–1.3 mm; valves torulose, glabrous; style 2–4 mm. Seeds
brown, oblong, 1.2–1.5 × 0.6–0.9 mm, wingless.
Habitat: Open gravel along streams, forests and clearings.
Elevation: 2100–4000 m.
Distribution: Bhutan, China (Sichuan, Yunnan).
Cardamine densiflora
Cardamine dentata
Cardamine Dep. ressa J. D. Hooker, Fl. Antarct. 1: 6. 1844. TYPE: New Zealand, moist places
near the sea at Rendezvous Harbour, J. D. Hooker s.n. (holotype, K).
Cardamine digitata Richardson, Bot. App. Frankl. Narr. Journey Polar Sea 743. 1823, not C.
digitata (Lamarck) O. E. Schulz (1903). TYPE:
Cardamine digitata var. oxyphylla Tratvetter; C. hyperborea var. oxyphylla (Trautvetter) O. E.
Schulz, Bot. Jahrb. Syst. 32: xxx. 1903. TYPE:
C. hyperborea O. E. Schulz, Bot. Jahrb. Syst. 32: 550. 1903. TYPE:
C. richardsonii Hultén, Acta Univ. Lund. N. F., Adv. 2, 41 (Fl. Alaska & Yukon): 838. 1945.
TYPE:
Herbs, perennial, glabrous throughout. Rhizomes slender, cylindrical, 0.5–1.5 mm in diam.;
stolons absent. Stems (0.6–)1–2(–3) dm, erect, simple, glabrous. Rhizomal leaves pinnately 5–7foliolate, not fleshy, (2–)4–10 cm; petiole (1.5–)3–7(–9) cm; terminal leaflet linear to narrowly
oblong or narrowly lanceolate, (0.5–)1–2.5(–3.3) cm × 0.5–4(–8) mm, sessile, glabrous, base
cuneate to attenuate, margin entire; lateral leaflets about as large as or smaller than terminal
leaflet, similar in other aspects; cauline leaves (1 or)2 or 3, 5–7-foliotate, alternate; petiole (2–)5–
12(–15) mm, base not auriculate; terminal leaflet narrowly lanceolate to linear, 1–3.2(–4) cm ×
0.5–2.5(–3.5) mm, sessile, base cuneate to attenuate, margin entire, apex acute to acuminate;
lateral leaflets similar to terminal one but smaller, sessile. Racemes ebracteate; fruiting pedicels
suberect to divaricate-ascending, (0.7–)1–2.5 cm. Sepals ovate, 2.5–3.5 × 1.5–2.5 mm, erect,
base of lateral pair not saccate; petals white, obovate, 5–9 × 2.5–5 mm, clawed, apex rounded;
37
median filament pairs 3–4.5 mm, lateral pair 1.5–2.5 mm; anthers oblong, 1–1.5 mm. Fruit
linear, (1.5–)2–4 cm × 1.5–2 mm; valves glabrous; style (1–)1.5–2.5(–4) mm; ovules and seeds
6–12 per fruit. Seeds brown, oblong, 1.5–2.2 × 0.9–1.1 mm. 2n = 28, 42.
Flowering: Jun–Aug.
Habitat: damp flats, river banks, tundra, meadows, bluffs, hummocks, sandy beaches, slopes,
mossy mats, sedge swales.
Elevation: 0 –1400 m.
Distribution: Canada (Northwest Territories, Yukon), Russia (Siberia, Far East), United States
(Alaska).
Cardamine diphylla (Michaux) A. Wood, Amer. Bot. & Fl. 37. 1870; Dentaria diphylla
Michaux, Fl. Bor.-Amer. 2: 30. 1803. TYPE:
Dentaria bifolia Stokes, Bot. Mat. Med. 3: 443. 1812. TYPE:
Dentaria incisa Small, Fl. SE U.S. 480. 1903, not D. incisa Eames, Rhodora 5: 216. 1903.
TYPE:
Herbs, perennial, glabrous or rarely sparsely pubescent. Rhizomes fleshy, cylindrical, 2–10
mm in diam., unsegmented, not fragile, with dentate leaf scars, somewhat uniform in diam.;
stolons absent. Stems (1.2–)1.5–3.5(–4) dm, erect, simple, glabrous or rarely sparsely pubescent
distally. Rhizomal leaves 3-foliolate, (5.5–)8–22(–26) cm; petiole (3–)4.5–16(–20) cm; terminal
leaflet ovate-elliptic to broadly ovate, (2–)3.5–8(–10) × (0.5–)2–6.5(–8) cm, with a petiolule 0.5–
1.2 cm, base cuneate to obtuse, margin coarsely crenate or dentate, puberulent with trichomes to
0.1 mm; lateral leaflets often similar to terminal one in shape, size and margin, base often
oblique, subsessile or petiolulate; cauline leaves 2(or 3), opposite or nearly so, 3-foliolate,
resembling rhizomal leaves; petiole (1–)2–4.5 cm, base not auriculate; terminal leaflet broadly
elliptic to ovate, (2–)4–8(–10) × 1–5 cm, coarsely dentate or crenate, minutely puberulent along
margin, subsessile or with petiolule 2–10 mm; lateral leaflets similar to terminal one. Racemes
ebracteate; fruiting pedicels ascending to divaricate, (1–)1.5–3(–3.6) cm. Sepals oblong, (4–)5–8
× 2–3 mm, erect, base of lateral pair slightly saccate; petals white or pink to purple, obovate to
oblanceolate, (7–)9–15(–17) × (3–)4–7 mm, short clawed, apex rounded; median filament pairs
5–8 mm, lateral pair 3.5–6 mm; anthers linear to oblong, 2.5–3 mm. Fruit linear-lanceolate, 1.5–
4 cm × 1.5–2.5 mm; valves glabrous; style 4–8(–10) mm; ovules and seeds 10–14 per fruit.
Seeds rarely produced, brown, oblong, 2–2.2 × 1.2–1.5 mm; cotyledons incumbent. 2n = 96.
Flowering: Apr–Jun.
Habitat: wooded bottoms and ravines, cliffs, bluffs, leges, shaded slopes, meadows, moist fields,
alluvial banks, rich woods.
Elevation: 50–1300 m.
Distribution: Canada (New Brunswick, Nova Scotia, Ontario, Quebec), United States (Alabama,
Arkansas, Connecticut, Georgia, Illinois, Indiana, Kentucky, Maine, Massachusetts, Michigan,
Minnesota, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania,
South Carolina, Tennessee, Vermont, Virginia, West Virgina, Wisconsin).
Cardamine dissecta (Leavenworth) Al-Shehbaz, J. Arnold Arb. 69: 82. 1988; Dentaria dissecta
Leavenworth, Amer. J. Sci. 7: 62. 1824. TYPE: United States, Alabama, Cherokee Co.,
Leavenworth s.n. (holotype, ).
Dentaria multifida Muhlenberg ex Elliott, Sketch Bot. S. Carolina 2: 142. 1822; C. multifida
(Muhlenberg ex Elliott) A. Wood, Amer. Bot. & Fl. 38. 1870, not C. multifida Pursh, Fl.
Amer. Sept. 2: 440. 1814; Cardamine angustata O. E. Schulz var. multifida (Muhlenberg ex
Elliott) H. E. Ahles; C. laciniata (Muhlenberg ex Willdenow) A. Wood subsp. multifida
38
(Muhlenberg ex Elliott) O. E. Schulz; C. laciniata var. multifida (Muhlenberg ex Elliott) J. F.
James, Bot. Gaz. 8: 207. 1883; D. heterophylla Nuttall var. multifida (Muhlenberg ex Elliott)
H. E. Ahles, J. Elisha Mitchell Sci. Soc. 80: 172. 1964; D. laciniata Muhlenberg ex
Willdenow var. multifida (Muhlenberg ex Elliott) J. F. James, Bot. Gaz. 13: 234. 1888.
TYPE: United States, mountaine of Carolina, Schweinitz s.n. (holotype, CHARL).
Dentaria furcata Small, Fl. SE U.S. 480. 1903. TYPE: United States, Alabama, near Tuscaloosa,
Rev. W. Johnson s.n. (holotype, NY!).
Herbs, perennial, glabrous throughot. Rhizomes fleshy, segments oblong, moniliform, 3–5
mm in diam.; stolons absent. Stems 1–3.5 dm, erect, simple, glabrous. Rhizomal leaves
palmately compound, biternate, 7–20 cm; petiole 4–13 cm; terminal leaflet ternately or pinnately
lobed, petiolule 1–3(–4) cm; ultimate segments linear, 0.7–3.5 cm × 0.6–3 mm, base attenuate to
cuneate, margin entire, apex apiculate; lateral leaflets about as large as or smaller than terminal
leaflet, similar in other aspects; cauline leaves (2 or) 3, palmately compound, biternate and
resembling rhizomal leaves, alternate to subopposite; petiole (0.5–)1–4(–5.5) cm, base not
auriculate; terminal leaflet ternate, petiolule (0.3–)0.7–1.5(–2.5) cm; ultimate segments linear,
(0.4–)1.5–5(–6.5) cm × 0.7–4(–6) mm, base cuneate to attenuate, margin entire; lateral leaflets
similar to terminal one. Racemes ebracteate; fruiting pedicels ascending to divaricate, 1–2.5(–
3.5) cm. Sepals oblong, 4–6 × 1.5–2 mm, erect, base of lateral pair not saccate; petals white to
pale pink, oblanceolate, 9–15 × 2–4 mm, not clawed, apex rounded; median filament pairs 6–7.5
mm, lateral pair 3–5 mm; anthers linear, 1.5–2.5 mm. Fruit linear-lanceolate, 2–3.5 cm × 1.5–2
mm; valves glabrous; style 4–7(–10) mm; ovules and seeds 10–14 per fruit. Seeds brown,
oblong, 1.5–2.5 × 1–1.5 mm. 2n = 64, 112.
Flowering: Mar–May.
Habitat: oak-hickory woods, moist loamy areas, floodplain woods, bluffs, rocky calcareous
woods, limestone slopes, along streams.
Elevation: 0–300 m.
Distribution: United States (Alabama, Georgia, Indiana, Kentucky, Ohio, North Carolina,
Tennessee).
Cardamine douglassii Britton, Trans. New York Acad. Sci. 9: 8. 1889 Dentaria douglassii
(Britton) Greene, Pittonia 3: 124. 1896. TYPE:
Arabis rhomboidea Persoon var. purpurea Torrey, Amer. J. Sci. 4: 66. 1822; Dracamine
purpurea (Torrey) Nieuwland, Amer. Midl. Naturalist 4: 40. 1915. TYPE:
Cardamine douglassii var. albidula Farwell, Amer. Midl. Naturalist 9: 26. 1925. TYPE:
Thlaspi tuberosum Nuttall, xxxxxxxx. TYPE:
Herbs, perennial, hirsute throughout or glabrous proximally. Rhizomes fleshy, tuberous at
stem base, subglobose, lobed or not, (3–)4–10 mm in diam.; stolons absent. Stems (0.7–)1–2.5(–
3) dm, erect, simple, sparsely to densely hirsute with trichomes (0.2–)0.3–0.6(–0.8) mm, or
glabrous near base. Rhizomal leaves simple, (3–)5–15(–18) cm; petiole (2–)4–12(–16) cm; blade
orbicular to cordate, sometimes reniform or ovate, (1–)2–6 × (0.7–)1.7–5 cm, base obtuse to
cordate, margin repand or entire; cauline leaves 3–6(–8), simple, not auriculate at base, middle
ones short petiolate, 2–5 × 0.5–2.5 cm, upper leaves sessile, oblong to ovate or lanceolate, entire,
repand, or coarsely dentate. Racemes ebracteate; fruiting pedicels ascending to divaricate,
(1–)1.5–3.5(–5) cm, sparsely pubescent or glabrous. Sepals oblong, 2.5–4(–6) × 1.5–2.5 mm,
erect, often hirsute, base of lateral pair not saccate; petals rose-purple to pink of very rarely
white, obovate, (7–)8–13(–15) × 3–5 mm, short clawed, apex rounded; median filament pairs 4–
7 mm, lateral pair 2–4 mm; anthers oblong, 1.3–1.7 mm. Fruit linear, (1.5–)2–4 cm × 1.5–2 mm;
39
valves glabrous; style 2–5 mm; ovules and seeds 10–16 per fruit. Seeds brown, oblong to ovate,
1.7–2.5 mm × 1–5 mm. 2n = 56, 64, 96, 112, 144.
Flowering: Mar–May.
Habitat: rich woods, bluffs, mesic bottomland foress, rocky hillsides, floodplains, seepage of
bogs, springy areas.
Elevation: 50–400 m.
Distribution: Canada (S Ontario), United States (Alabama, Connecticut, District of Columbia,
Illinois, Indiana, Iowa, Kentucky, Maryland, Massachusetts, Michigan, Minnesota, Missouri,
New Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee,
Virginia, West Virgina, Wisconsin).
Cardamine elegantula J. D. Hooker & Thomson, J. Linn. Soc., Bot. 5: 146. 1861. TYPE:
Bhutan, 6500 m, Griffith s.n. (holotype, K!; isotype, P!).
Herbs, perennial, slender, pilose on leaves and stems with trichomes to 0.6 mm. Rhizomes
slender, stoloniferous. Stems 8–30 cm, simple, flexuous. Basal leaves compound, rosulate, often
present by fruiting; petiole 0.7–2 cm; terminal leaflet broadly ovate to obovate, petiolulate,
sparsely pilose, base distinctly oblique, margin entire or obscurely 1-toothed on each side; lateral
leaflets 2–4 on each side, with a petiolule 0.5–1.5 mm, ovate to elliptic, 2–5(–8) × 1–3.5(–5) mm,
similar to terminal one or with a lateral lobe on abaxial margin, distinctly mucronate; cauline
leaves compound, not auriculate at base; petiolule 0.5–3 mm; terminal leaflet elliptic to linearelliptic, 3–6(–9) × 0.5–2 mm, base attenuate to cuneate and decurrent with adjacent lateral lobes,
margin entire and scabrous with trichomes 0.05–0.1 mm, apex mucronate; lateral leaflets 5–8 on
each side of rachis, petiolulate, similar to terminal leaflets. Racemes ebracteate, 2–5-flowered;
rachis strongly flexuous; fruiting pedicels strongly reflexed, slender, straight, 4–6 mm, glabrous.
Sepals ovate, 1.5–2 × 0.8–1 mm, glabrous, membranous at margin and apex; petals pink or
white, broadly obovate, 6–7 × 3–4 mm; median filament pairs 2.5–3 mm, filiform; lateral pair
1.5–2 mm; anthers oblong, 0.4–0.5 mm; ovules 14–20 per ovary. Fruit linear, 1.2–1.5 cm × 0.7–
1 mm; valves glabrous; style 1–2 mm. Seeds pale brown, oblong, ca. 1.8 × 0.8 mm, wingless.
Flowering: May–Jun.
Habitat: beneath wet rock overhang, damp rocks in forests, marshy area along streams.
Elevation: 3050–2400 m.
Distribution: Bhutan, India (Sikkim), Nepal.
Cardamine engleriana O. E. Schulz, Bot. Jahrb. Syst. 32: 407. 1903. TYPE:
Cardamine griffithii J. D. Hooker & Thomson var. grandifolia T. Y. Cheo & R. C. Fang, Bull.
Bot. Lab. North-East. Forest. Inst., Harbin 1980(6): 25. 1980. TYPE: China, Hubei, Xingshan
Xian, 1,600 m, 27 May 1957, Y. Liu 529 (holotype, NAS).
Herbs, perennial, sparsely to densely pilose, often glabrescent above, rarely glabrous
throughout. Rhizomes slender, with 1 or few filiform stolons. Stems 10–24(–30) cm, erect,
simple, terete, crisped pilose at base. Basal leaves not rosulate, often withered by anthesis;
petiole 0.5–2 cm; terminal leaflet reniform or suborbicular, 0.5–1.5 × 0.7–1.7 cm, base cordate,
margin shallowly crenate or repand; lateral leaflets 2, much smaller than terminal ones,
sometimes absent and leaf simple; middle and upper cauline leaves trifoliolate, sessile, to 10 cm;
terminal leaflet reniform, suborbicular, or broadly ovate, 1.5–6(–7) × 1.2–4(–5) cm, with a
petiolule 0.3–2.5(–5.2) cm, base cordate, rounded, or cuneate, glabrous, ciliate, or trichomes
forming small tufts at teeth, margin crenate, repand, or obscurely to distinctly 2–6-toothed on
each side, apex obtuse or acute; lateral leaflets 2, auriclelike, attached at or just above node, often
40
giving appearance of amplexicaul leaf base, suborbicular, ovate, obovate, or reniform, 3–16(–25)
× 2–13(–17) mm, much smaller than terminal leaflet, margin entire, 1- or 2-toothed, or repand.
Racemes ebracteate, 3–10-flowered; fruiting pedicels 0.5–1.7 cm, divaricate or ascending,
slender. Sepals ovate, 2.5–3 × 1–1.5 mm, lateral pair subsaccate; petals white, obovate, 6–8 × 2–
3 mm, apex rounded; median filament pairs 3–4.5 mm; lateral pair 1.5–3 mm; anther oblong,
0.7–1 mm; ovules 8–16 per ovary. Fruit linear, slightly flattened, 1.5–2.5 cm × 1–1.2 mm;
valves smooth, glabrous; style 1–2 mm. Seeds brown, oblong, 1.5–1.8 × 0.8–1 mm, wingless.
Flowering: Apr−Jul.
Habitat: shady slopes, woodlands, moist places in ravines.
Elevation: 800–2900 m.
Distribution: China (Anhui, Gansu, Hubei, Shaanxi, Sichuan).
Cardamine enneaphyllos (Linnaeus) Crantz, Cl. Crucif. Emend. 127. 1769; Dentaria
enneaphyllos Linnaeus, Sp. Pl. 2: 653. 1753. TYPE: “Dentaria triphyllos Bauhin. As salinas
Austriae Superioris, Herb. Burser no. XVIII(1): 83 (lectotype designated by Marhold (2001:
44), UPS).
Cardamine enriquei Marhold, Lihová & Perný, Bot. J. Linn. Soc. 139: 293. 2002. TYPE:
Cardamine eremita Standley & Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23: 53. 1944; C.
bonariensis Persoon subsp. eremita (Standley & Steyermark) Sjöstedt, Bot. Not. 128: 15.
1975. TYPE: Guatemala, J. Steyermark 50143 (holotype, F!; isotype, US!).
Herbs, perennial, cespitose, with slender caudices, glabrous throughout. Rhizomes absent.
Stems 8–20 cm, erect or ascending, simple or branched above. Basal leaves and lowermost
cauline 2-4 cm, petiolate, not auriculate; terminal leaflet 3–7(–10) × 2–5 mm, linear-oblanceolate
to oblanceolate, sessile, base attenuate, margin entire, apex obtuse; lateral leaflets 2–4 on each
side of rachis, similar to terminal leaflet but smaller; uppermost leaves smaller, becoming simple.
Racemes bracteate basally, secund and dense in fruit; fruiting pedicels 5–9 mm, ascending,
slender, straight. Sepals 1.5–2 × 0.7–1 mm, oblong, caducous; petals 3.5–5 × 1–1.5 mm, white,
spatulate, apex obtuse; filaments 2–2.5 mm; anthers 0.2–0.3 mm; ovules 12–14 per ovary. Fruits
2–2.7 cm × 0.8–0.9 mm; style 1–1.8 mm. Seeds 1.5–1.7 × ca. 0.5 mm, brown, oblong.
Flowering: Feb–Aug.
Habitat: rocky areas in alpine pine forests, limestone ridges, meadows.
Elevation: 2800–3700 m. (Endemic).
Distribution: Guatemala.
Cardamine fallax (O. E. Schulz) Nakai, Rep. Veg. Ooryongto 19. 1919; C. flexuosa subsp.
fallax O. E. Schulz, Bot. Jahrb. Syst. 32: 478. 1903; C. flexuosa var. fallax (O. E. Schulz)
Nakai, J. Coll. Sci. Imp. University Tokyo 31: 441. 1911; C. flexuosa var. fallax (O. E.
Schulz) T. Y. Cheo & Fang, Bull. Bot. Lab. N.E. Forest. Inst. Harbin 1(6): 23. 1980, isonym;
C. scutata subsp. fallax (O. E. Schulz) H. Hara, J. Fac. Sci. Univ. Tokyo, sect. 3, Bo. 6(2): 59.
1952. TYPE: [Japan], Mama-mura, Shimosa, anonymous, 11 May 1894 (lectotype designated
by Marhold et al. (2007: 657), US00324475!).
Cardamine fargesiana Al-Shehbaz, Novon 10: 324. 2000. TYPE: China, Sichuan, Tchen-kéoutin, R. P. Farges 1341 bis (holotype, P!; isotype, P!).
41
Herbs, perennial. Rhizomes ovoid or oblong, 2–5 × 1.5–2 mm; stolons slender, densely
pilose with crisped trichomes. Stems 8–23 cm. simple, erect, densely crisped pilose at base,
gradually sparser upward, completely glabrous at raceme. Basal leaves 1–4 per plant, simple or
rarely with a pair of leafletlike lobes; petiole 0.5–4.5 cm, densely crisped pilose; leaf blade
orbicular, 5–12 mm in diam., palmately 5-veined, abaxially glabrous or sparsely pilose along
proximal portion of veins, adaxially pilose with simple, straight trichomes to 0.5 mm, base
cordate, margin entire or obscurely 5-angled, mucronate at vein tips; cauline leaves 3–5, 0.9–2.2
× 0.7–1.5 cm, broadly ovate or oblong in outline, pinnatisect, abaxially glabrous, adaxially
pilose; petiole absent and leaf base appearing auriculate; rachis 2–15 mm; terminal lobe of
lowermost leaf obovate, 4–14 × 2–6 mm, apically 3-toothed, sometimes with a minute tooth
subbasally on each side, base cuneate into a petiolule to 3 mm, margin entire, apex mucronate;
lateral lobes 4–6, linear to lanceolate-linear, margin entire, apex acute, proximal 2–4 attached at
one point on each side of rachis. Racemes ebracteate, 3–7-flowered; flowering pedicels
ascending, 5–12 mm, straight, glabrous. Sepals ovate, 2.5–3 × 1.2–1.6 mm, glabrous, margin and
apex membranous, lateral pair saccate; petals white, broadly obovate, 5–6 × 2.5–3 mm, apex
rounded; median filament pairs 3–3.5 mm, slender, lateral pair 2–2.5 mm; anthers oblong, 0.8–
1.1 mm, base sagittate. Fruit (young) glabrous; style 1–1.5 mm; stigma entire. Seeds not seen.
Distribution: China (Sichuan).
Notes: Known only from two collections bothy made in late 19th century.
Cardamine flagellifera O. E. Schulz, Bot. Jahrb. Syst. 32: 405. 1903. TYPE:
Cardamine hugeri Small, Man. Fl. SE. U.S. 569. 1933; Crdamine flagellifera var. hugeri (Small)
Rollins, Cruciferae Continental N. Amer. 274. 1993. TYPE:
Herbs, perennial, hirsute below, sparsely pubescent or glabrous above. Rhizomes fleshy,
swollen and compact at stem base, producing many, slender, pubescent or glabrous stolons ca. 1
mm in diam. Stems 1–2.5 dm, erect, simple or rarely few branched above, sparsely to densely
hirsute or pilose near base, sparsely so or glabrous distally. Rhizomal leaves simple or rarely 3foliolate, (3–)6–16 cm; petiole (1.5–)4–13 cm; blade or terminal leaflet orbicular to reniform or
broadly ovate, (1–)1.5–4.5 cm, petiolule 0.5–2 cm, base cordate, margin repand to coarsely
crenate and apiculate at vein endings, sparsely pubescent or glabrous, not puberulent; lateral
leaflets much smaller than terminal one, subsessile or on petiolules to 0.5 cm; cauline leaves 2–5,
3–5-foliotate; petiole 0.8–5 cm, base not auriculate; terminal lobe broadly ovate to suborbicular,
1.5–4(–5) × 1.2–3.5 cm, repand to coarsely creanate or slightly lobed, apiculate at vein endings,
glabrous along margin, petiolule 0.5–1.5 cm; lateral leaflets similar to terminal but smaller,
sessile or on petiolule to 0.5 cm. Racemes ebracteate; fruiting pedicels ascending to divaricateascending, 0.7–2 cm. Sepals oblong, 3–3.5 × 1–1.5 mm, ascending, base of lateral pair not
saccate; petals white, oblanceolate, 5–6.5(–8) × 1.5–2.5 mm, not clawed, apex obtuse to
subemarginate; median filament pairs 3.5–5 mm, lateral pair 2.5–3 mm; anthers oblong, 1.2–1.6
mm. Fruit linear, 1.5–2.5 cm × 1–1.2 mm; valves glabrous; style 1.2–2.5 mm; ovules and seeds
10–14 per fruit. Seeds brown, oblong, 1.2–1.7 × 0.8–1.1 mm.
Flowering: Mar–Jun.
Habitat: moist wooded slopes, gorges, wooded ravines, seepage places.
Elevation: 300–1000 m.
Distribution: United States (Georgia, North Carolina, South Carolina, Tennessee, West Virgina).
Cardamine flexuosa Withering, Arr. Brit. Pl., ed. 3, 3: 578. 1796; C. hirsuta Linnaeus subsp.
flexuosa (Withering) Forbes & Hemsl., J. Linn. Soc., Bot. 23: 43. 1886; C. hirsuta Linnaeus
42
var. flexuosa (Withering) J. D. Hooker & T. Anderson in J. D. Hooker, Fl. Brit. India 1: 138.
1872; C. scutata Thunberg subsp. flexuosa (Withering) H. Hara, J. Fac. Sci. Univ. Tokyo III,
6: 59. 1952. TYPE: “Rookery at Edgebaston”. Lectotype (designated by Post et al. [J. Bot.
Res. Inst. Texas 3(1): 227. 2009]): Curtis Flora Londinensis t. 277. 1777.
Cardamine arisanensis Hayata, Icon. Pl. Formos. 3: 20. 1912; Barbarea arisanensis (Hayata) S.
S. Ying, Alp. Pl. Taiwan Color. 2: 170. 1978. TYPE:
Cardamine debilis D. Don, Prodr. Fl. Nepal. 201. 1825, not Banks ex de Candolle, Syst. Nat. 2:
265. 1821; Cardamine flexuosa Withering subsp. debilis O. E. Schulz, Bot. Jahrb. Syst. 32:
478. 1903; Cardamine flexuosa Withering var. debilis (O. E. Schulz) T. Y. Cheo & R. C.
Fang, Bull. Bot. Lab. North-East. Forest. Inst., Habrin (1980)6: 23. 1980. TYPE:
Cardamine sylvatica Link var. kamtschatica Regel, Bull. Soc. Imp. Naturalistes Moscou 34: 172.
1861; C. flexuosa Withering var. kamtschatica (Regel) Matsumura, Bot. Mag. (Tokyo) 13: 73.
1899. TYPE:
Cardamine flexuosa Withering var. ovatifolia T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. NorthEast. Forest. Inst., Habrin (1980)6: 24. 1980. TYPE: China. Zhejiang: Hangchow Longjing,
21 March 1957, S. Y. Chang 186 (holotype, NAS).
Cardamine hirsuta Linnaeus var. flaccida Franchet, Bull. Soc. Bot. France 33: 398. 1886. TYPE:
China. Yunnan, Mosoyn, near Lankong, 14 March 1885, Delavay 1839 (holotype, P!; isotype,
US!).
Cardamine hirsuta Linnaeus var. omeiensis T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. NorthEast. Forest. Inst., Habrin 6: 23. 1980. TYPE: China. Sichuan: Mount Omei, Jinding, 1 July
1938, 3035 m, H. C. Chou 7690 (holotype, WH (as HWH); isotypes, A!, NAS!).
Cardamine occulata Horneman, Hort. Hafn. Suppl. 71. 1919; C. flexuosa Withering subsp.
debilis var. occulata (Horneman) O. E. Schulz, Bot. Jahrb. Syst. 32: 479. 1903. TYPE:
Cardamine sylvatica Link in Hoffmann, Phyt. Blätt. 1: 50. 1803. TYPE:
Cardamine zollingeri Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 28: 294. 1854.
Nasturtium obliquum Zollinger & Mor., Nat. en Geneesk. Arch. Neerl. Indie 2: 580. 1845.
TYPE:
Herbs, annual or biennial, sparsely to densely hirsute basally or throughout, or glabrous. Stem
(6–)10–50 cm, erect, ascending, or decumbent, 1 to several from base, simple or branched,
flexuous or straight. Basal leaves not rosulate, often withered by anthesis, petiolate; leaf blade
(2–)4–10(–14) cm, lyrate; terminal lobe reniform, broadly ovate, or suborbicular, repand or 3–5lobed; lateral lobes, (1 or)2–6(or 7) on each side of midvein, petiolulate or subsessile, oblong,
ovate, or elliptic, smaller than terminal lobe, entire, repand, crenate, or 3(–5)-lobed; cauline
leaves 3–15, including petiole (2–)3.5–5.5(–7) cm; petiole base not auriculate; terminal lobe 3–5lobed; lateral lobes 2–7 on each side of midvein, suborbicular, ovate, oblong, oblanceolate, or
linear, similar to or slightly smaller than terminal lobe, sessile or shortly petiolulate, entire,
repand, dentate, or 3(–5)-lobed. Raceme ebracteate, several flowered; fruiting pedicels divaricate
or ascending, (5–)6–14(–17) mm, slender. Sepals oblong, 1.5–2.5 × 0.7–1 mm; petals white,
spatulate, 2.5–4(–5) × 1–1.7 mm; stamens 6, rarely 4 and lateral pair absent; filaments 2–3 mm;
anthers ovate, 0.3–0.5 mm; ovules 18–40 per ovary. Fruit linear, (0.8–)1.2–2.8 cm × 1–1.5 mm;
valves glabrous, torulose; style 0.3–1(–1.5) mm. Seeds brown, oblong or subquadrate, 0.9–1.5 ×
0.6–1 mm, narrowly margined or not. 2n = 32.
Flowering: Feb–Jun.
Habitat: Fields, roadsides, grasslands, disturbed sites, streamsides, clearings, running water, wet
forests, dry sites, greenhouses.
Elevation: 0–3600 m.
43
Distribution: native in Europe (all countries except Iceland); naturalized in Africa (Algeria,
Morocco), Asia [Bangladesh, Bhutan, China (Guangdong, Guizhou, Sichuan, Yunnan), India
(Sikkim), Indonesia, Japan, Kashmir, Korea, Laos, Malaysia, Myanmar, Nepal, Pakistan,
Philippines, Taiwan, Thailand, Vietnam], naturalized in Australia, North America [Canada
(British Columbia, Newfoundland, Ontario), United States (Alabama, California, Florida,
Georgia, Illinois, Indiana, Louisiana, Maryland, Michigan, New York, North Carolina, Ohio,
Oregon, Rhode Island, Texas, Virginia, Washington)], Central America (Mexico), and South
America [Argentina (Corrientes, Jujuy, Salta), Bolivia (La Paz, Santa Cruz), Brazil (LinharesEspirito Santo), Colombia (Antioquia), Paraguay, Peru (Cajamarca, Pasco), Venezuela
Specimens examined: ARGENTINA. Corrientes: San Cosme, 16 Dec. 1992, Ishikawa s.n.
(MO); Corrientes, Krapovickas 39348 (BACP, LIL), Schinini 35131 (NY). Jujuy: Capital,
Rotman 691 (SI). Salta: San Martín, camino a Cambuco, 17 July 1980, Azcue s.n. (BACP).
BOLIVIA. La Paz: Sud Yungas, 2 km N Chulumani or rd to Irupana, 16°23′N, 67°32′W,
Solomon & Nee 14278 (MO). Santa Cruz: Andres Ibañez, Santa cruz de la Sierra, Irala Avenue
565, 17°46′S, 63°11′W, Abbott 16866 (MO). BRAZIL. Linhares-Espírito Santo: Reserve
Florestal de Linhares, Folli 3072 (MO); Cuitiba, Guaratuba, Hatschbach 43877 (BACP).
COLOMBIA. Antioquia: Titiribí, Verda “Corcovado,” sitio alto de Buenavista, 6°4′N,
75°48′W, González 19 (MO); Angelópolis, Gutiérres & Barcley 17c634 (GH). PARAGUAY.
Central, Asunción, Schinini 4132 (BAA, BACP). Valle. Cali, Can Antonio o Cerro La Horqueta,
Silverston-Sopkin 1490 (MO). PERU. Cajamarca: San Ignacio, San José de Lourdes, between
Camaná and Santo Tomás, 5°1′S, 78°52′W, Campos & Corrales 3798 (MO, NY); San José de
Lourdes Bosque secundario, Campos & Vásquez 6476 (MO). Pasco: Oxapampa, Abra de Villa
Rica, 10°40′S, 75°18′W, Rojas et al. 3260 (MO). VENEZUELA. Bolivar: Gran Sabana, Sierra
de Lema, La Escalera, 6°0′N, 61°23′W, Diaz et al. 6832 (MO). Distrito Federal: Parque
Nacional El Avila, Cordiller de la Costa, 10°34′N, 66°54′W, Meier 7770 (MO). Miranda: Los
Teques, Carretera hacia Laguneta, Montes 346 (S). Trujillo: Dtto. Bocono, Laguna Aguas
negras, 3 km SW of Batatal, 9°19′N, 70°11′W, Aymard & Stergios 2405 (NY, PORT).
(Bolivar, Distrito Federal, Miranda, Trujillo)].
Cardamine flexuosa is one of the most variable species of Brassicaceae, especially in plant
height, flower size, stem direction, density of indumentum, orientation of fruiting pedicels,
degree of zigzagging of the raceme rachis, and, most dramatically, shape, number, size, margin,
and base (sessile vs. petiolulate) of lateral leaf lobes. Although Schulz (Bot. Jahrb. Syst. 32: 464–
473. 1903) recognized 20 infraspecific taxa in C. flexuosa, an examination of the types of most
of these reveals that he confused the limits of least four different species, including C. parviflora,
C. scutata, and the North American C. pensylvanica Muhlenberg ex Willdenow. Only a small
fraction of the vast number of variants of C. flexuosa have been accorded infraspecific ranks, but
all of those occur sporadically throughout the species range, and we prefer to avoid the
recognition of infraspecific taxa that are based on unreliable characters, especially the
morphology of lateral leaf lobes. Although C. occulata was based on Chinese material, it was not
mentioned in FRPS, and it represents only a minor variant of C. flexuosa.
Cardamine fragariifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 446. 1903. TYPE: China, Hupeh, A.
Henry 5803 (holotype, B; isotype, GH!).
Cardamine scoriarum W. W. Smith; Cochlearia scoriarum (W. W. Smith) Handel-Mazzetti.
TYPE: China, Yunnan, flank of volcanic mountain NW of Lengyueh?, 25°70′N, 7000 ft, Jun
1912, G. Forrest 8201 (holotype, B!).
44
Cardamine smithiana Biswas. TYPE: Donga La, 9500 feet, 25 Jul 1933, F. Ludlow & G. Sherriff
324 (holotype, BM!).
Cochlearia alatipes Handel-Mazzetti; Hilliella alatipes (Handel-Mazzetti) Y. H. Zhang & H. W.
Li; Yinshania alatipes (Handel-Mazzetti) Y. Z. Zhao. TYPE: China, Hunan, in monte
Yünschan, near Wukang. ca. 1000 m, 12 Jun 1918, H. F. Handel-Mazzetti 12,098 (holotype,
WU!; isotypes, E!, W!, WU!).
Herbs, perennial. Rhizomes 3–8 mm in diam. Stems (3.5–)5–10(–13) dm, erect, simple or
branched above, glabrous or rarely puberulent. Basal leaves withered by flowering; cauline
leaves (5–)7–10(–15), petiolate, trifolioate, rarely lowermost with 4 or 5 leaflets, often simple
below raceme; petiole 1–6(–10) cm, not auriculate at base, wingless or wing rarely to 1 mm
wide; petiolules 1–6(–11) mm; leaflets subequal or terminal larger, ovate or lanceolate, rarely
elliptic or subrhomboid, (1.5–)3–8(–12) × (1–)2–3(–5) cm, glabrous or with appressed, antrorse,
simple trichomes 0.1–0.7 mm, base cuneate or obtuse, that of lateral leaflets often oblique,
margin serrate or crenate, rarely lobed or subentire, ciliate with trichomes more than 0.1 mm,
teeth mucronate, apex acute, acuminate, or acuminate-caudate. Raceme ebracteate, several
flowered; fruiting pedicels divaricate, 5–9(–13) mm, slender, straight, glabrous or sparsely
puberulent. Sepals oblong, 2.5–3.5(–4) × 1.5–2 mm, erect or ascending, not saccate, glabrous;
petals purple, lavender, pink, or rarely white, obovate or obovate-oblong, (6–)7–9(–10) ×
(2.5–)3–4 mm, rounded at apex; claw 1.5–2(–3) mm; median filament pairs 4–5 mm, lateral pair
3–4 mm; anthers (0.6–)0.7–1 mm. Fruit (young) glabrous; style slender, ca. 1 mm; stigma entire.
Seeds not seen.
Flowering: May−Jul.
Habitat: volcanic mountain slopes, shaded streamsides, wet forests, rocky places, moist
grasslands.
Elevation: 1000–3000 m.
Bhutan, China (Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan), India, Myanmar.
Cardamine franchetiana Diels, Notes Roy. Bot. Gard. Edinburgh 5: 205. 1912; Loxostemon
delavayi Franchet, Bull. Soc. Bot. France 33: 400. 1886, not Cardamine delavayi Franchet,
Bull. Soc. Bot. France 33: 397. 1886. TYPE: China, Yunnan, Likiang, 9 Jul 1884, Delavay
[35] (holotype, P!).
Loxostemon smithii O. E. Schulz, Acta Hort. Gothob. 1: 161. 1924; Loxostemon smithii O. E.
Schulz var. glabrescens O. E. Schulz, Acta Hort. Gothob. 1: 162. 1924. TYPE: China,
Sichuan, SE of Matang, 15 Sep 1922, 4800–5100 m, H. Smith 4445 (holotype, UPS!).
Herbs, perennial, hirsute or glabrescent. Rhizomes to 1 cm, with stolons and numerous
bulbils; bulbils whitish, fleshy scales ovoid or subglobose, with rudimentary apical appendages.
Stems (5–)10–20(–27) cm, simple, erect; underground proximal part whitish, slender or filiform,
glabrous; aboveground part green or purplish, stout, pilose or glabrous. Rhizomal leaves 1 or 2,
4–12(–18) cm; petiole 4–10 cm; terminal leaflet oblong, 4–10 × 2–6 mm, sessile or with a
petiolule to 2 mm; lateral leaflets 2–4 pairs, similar to terminal one; cauline leaves (1 or)2 or 3(or
4), 1.5–5 cm; petiole 0.4–4(–6) cm, base not auriculate; terminal leaflet oblong to oblong-elliptic,
4–11 × 1.5–3 mm, ciliate or glabrous, base cuneate, margin entire, apex mucronate; lateral
leaflets (2 or)3–6 pairs, similar to terminal one. Raceme ebracteate, 3–8-flowered; fruiting
pedicels divaricate-ascending to ascending, 4–13 mm, straight. Sepals ovate or suboblong, 2–3 ×
1.2–1.7 mm, margin membranous; petals white or rarely lavender, broadly obovate or
suborbicular, 6–9 × (3–)4–6 mm, base abruptly narrowed into a claw 1–2 mm, apex rounded;
median filament pairs 3–4 × 0.4–0.6 mm, flattened, not extended apically into a tooth; lateral pair
45
slender, 2–3 mm; anthers oblong, 0.6–0.9 mm; ovules 8–16 per ovary. Fruit linear, 1.5–2.5 cm ×
1.5–2 mm; valves smooth, glabrous; style 1–3 mm. Seeds brown, broadly oblong or ovate, 1.3–
1.6 × 0.9–1.2 mm, wingless.
Flowering: Jun–Jul.
Habitat: mountain slopes, deep valleys, rocky areas, scree, crevices of boulders, meadows, moist
pastures.
Elevation: 2300–4800 m.
Distribution: China (Qinghai, Xizang, Yunnan).
Cardamine franklinensis I. Thompson, Muelleria 9: 152. 1996. TYPE: Australia, Australian
Capital Territory, 3.2 km above Bendora on Mt. Franklin Road, 13 Nov 1953, C. W. E. Moore
2777 (holotype, NSW).
Herbs, perennial, glabrous throughout. Stems erect, to 30 cm tall, usually branched from base
and above. Basal leaves densely rosulate, thin, petiolate, to 15 cm, simple or pinnate; simple
leaves spatualte, crenate or entire; pinnate leaves 3–5-foliolate, terminal lobe elliptic to ovate,
lateral leaflets sessile, obovate, attenuate at base; cauline leaves 0–3, petiolate, not auriculate at
base, reduced in size upwards, lower ones similar to basal leaves, uppermost simple, pinnatifid or
entire. Raceme ebracteate, many flowered; fruiting pedicels divaricate-ascending, ca. 10 mm.
Sepals ovate, 2–3 mm; petals strongly differentiated into blade and claw, white or pink on the
outside, 4.5–6.5 mm; stamens 6. Fruits suberect, 2.5–3.5 cm × 1.5–1.8 mm; style 1–2 mm.
Seeds elliptic, 1.3–1.5 mm.
Flowering:
Habitat: subalpine woodlands, rocky scree slopes, among Poa tussocks.
Distribution: Australia (Australian Capital Territory, New South Wales, Tasmania, Victoria).
Cardamine fulcrata Greene, Pittonia 3: 155 (1897). TYPE: Mexico, Pringle s.n. (holotype,
ND-G). US has three sheets as Pringle 4989 marked as isotypes: check original publication.
Cardamine aschersoniana O. E. Schulz, Bot. Jahrb. Syst. 32: 410. 1903. TYPE: Venezuela,
Tovar, May 1836, Moritz 369 (lectotype, here designated, B!).
Cardamine bradeii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 328. 1923. TYPE: Costa
Rica, Barba, 22 Jan. 1909, A. C. Brade 2305 (holotype, B?).
Cardamine fulcrata var. scabra O. E. Schulz, Bot. Jahrb. Syst. 32: 411. 1903. TYPE: Venezuela,
Tovar, 2340−2670 m, 1854−1855, A. Fendler 23 (holotype, GH!; isotypes, K!, NY!).
Herbs, perennial. Rhizomes slender, not tuberous. Stems (1.5−)2.5−6(−8) dm, erect or
decumbent, simple or branched above, glabrous or puberulent. Cauline leaves (lower and
middle) 3.3−16(−23) cm, trifoliolate; petiole 2−8(−10) cm; terminal leaflet 2−8(−11) ×
(0.7−)1.5−3(−4) cm, lanceolate, oblong, elliptic, or ovate, sparsely hirsute or puberulent with
trichomes to 0.5 mm, base cuneate to subobtuse, margin ciliolate with trichomes to 0.2 mm,
serrate or crenate, teeth mucronate, apex acute to long acuminate, petiolule 0.5−1.5 cm; lateral
leaflets smaller than terminal lobe, subsessile or with a petiolule to 5 mm, base oblique;
uppermost leaves smaller, reduced into bracts. Racemes bracteate throughout or rarely only
basally; bracts simple, linear to linear-oblanceolate; lowermost filiform to lanceolate and to 5 ×
1.5 cm, subsessile or on petioles to 2 cm; fruiting pedicels (7−)10−20(−25) mm, divaricate to
ascending, stout. Sepals oblong, 2−3.5 × 1−1.5 mm, not saccate; petals white, obovate to
oblanceolate, 4.5−7 × 2−4 mm; filaments 2.5−4.5 mm; anthers oblong, 0.8−1.2 mm; ovules
46
12−20 per ovary. Fruits (2.8−)3.5−5(−5.8) cm × 1.5−2.2 mm; style 1−2.7(−4) mm. Seeds brown,
oblong, 1.5−2.7(−3) × 0.8−1.2(−1.5) mm.
Flowering: sporadically throughout the year.
Habitats: Cloud or rain forests, shady hillsides, mossy stream banks, along trails, forest margins.
Elevation: 700–3300 m.
Habitat: cloud forests, shady hillsides, mossy stream banks, along trails in forests, forest margins.
Elevation: 700–2800 m. Distribution: Southern Mexico (Chiapas), Central America (Costa Rica,
Guatemala, Honduras, Panama) and northern South America [Colombia (Magdalena,
Santeander), Venezuela (Lara, Merida, Monagas, Sucre, Tachira, Trujillo)].
Specimens examined: COLOMBIA. Magdalena: E of La Finca de Cortes, 0.5 km, 10°54′N,
73°58′W, Kirkbride & Forero 1860 (US). Santander: W slope of Cordillera Oriental, hwy from
Bucaramanga to Cúcuta, km 31, Stein 3622 (GH, MO, NY); Chitagá Valley, 1 km N Alto del
Almorzadero, 73°39-40′W 06°59′N, St. John 20794 (GH); Zapatoca, Fassett 25519 (GH).
VENEZUELA. Lara: Dtto. Moran, trail from Humocaro to Buenos Aires (Caserio) below
Páramo Los Rosas, Liesner et al. 8160 (MO). Merida: La Carbonera, Bernardi, Charpin &
Jacquemoud 17188 (F, G); San Rafael, Páramo de Las Coloradas, Cuatrecasas, Ruiz-Terán &
López-Figueiras 28521 (F, US); Andrés Bello, Zerpa, Wessels Boer 1737 (F, K, MO, NY);
Campo, Bosque de San Eusebio, Ruíz-Terán 1095 (MO); La Mucuy, 15 km NEE of Mérida,
Breteler 3474 (NY, S, US); La Mucuy, Rio Loro, on trial to La Lagtuna Coromoto, Barclay &
Juajibioy 9936 (GH, MO); trail from Cabin at La Escalera to the Puente de La Escalera, Luteyn
et al. 6213 (MO, NY); Mucuquí, Bosque del Cacique, Bernardi 177 (NY). Monagas: Cuácharo
Cave, 6 km NW Guácharo, Steyermark 62314 (F, GH). Sucre: Cerro Turumuquire, headwaters
of Río Manzanares, Steyrmark 62685 (F). Tachira: Páramo Zumbador, Charpin, Jacquemoud &
Ruíz-Terán 13392 (F, G); 3−8 kms de Zumbador, between Zumbador and Queniquea, Styermark
105081 (MO); Quebrada Agua Azul, S El Reposo, 14 km SE Delicias, 7°31′V, 72°24′W,
Steyermarck & Liesner 118343 (GH, MO). Trujillo: Boconó, Parque Nacional Guaramacal,
Laguna de Agua Negra, Qda Salvaje, Stergios, Dorr & Caracas 20295 (MO, US); sector
Quebrada Honda de El Santuario, 1012298N, 359185E, Stergios, Niño & Cuello 19338 (MO,
US); Guaramacal Paramo, Stergios & Caracas 19226 (MO, US); Dorr et al. 4980 (GH, NY).
The only difference used by Schulz (1903) to separate Cardamine fulcrata from C.
aschersoniana in having smaller and acute instead of lager and acuminate terminal leaflets of
cauline leaves. By contrast, the differences he (Schulz, 1923) used to separate C. fulcrata from
C. bradeii is having sessile instead of long petiolulate (to 5 mm Long) lateral leaflets. In my
opinion, these differences break down when the overall variability of the species in its entire
range is taken into consideration. Furthermore, Schulz (1903) cited one collection, Fendler 23
(GH) under both C. furcata and C. aschersoniana.
Cardamine fulcata is often confused with C. africana, an African species naturalized in the
Neotropics, but it is easily distinguished by having bracteate instead of ebracteate inflorescences.
Cardamine gallaecica (M. Laínz) Rivas Mart. & Izco, Itinera Geobot. 15: 699. 2002.
Cardamine georgiana Al-Shehbaz & Warwick, Rhodora 101: 275. 1999; Dentaria microphylla
Willdenow, Sp. Pl. 3: 479. 1800; C. microphylla (Willdenow) O. E. Schulz, Bot. Jahrb. Syst.
32: 342. 1903, not C. microphylla Adams, Mém. Soc. Imp. Naturalistes Moscou 5 : 111.
1817, nor C. microphylla J. Presl, Delic. Prag. 1 : 15. 1822. TYPE: Georgia, anonymous
(holotype, B-W! ).
47
Cardamine geraniifolia (Poiret) de Candolle, Syst. nat. 2: 268. 1821; Sisymbrium geraniifolium
Poiret in Lamarck, Encycl. 7: 218. 1806; Dentaria geraniifolia (Poiret) Reiche, Fl. Chile 1:
104. 1896. TYPE: Chile, Magellan, Commerson s.n. (holotype, P-JU! ; isotypes, 2P!)
Herbs, perennial. Rhizomes fleshy, elongated, non-scaly, to 12 cm × 6 mm. Stems
(14−)20−38(−46) cm, erect or decumbent only at base, simple or branched above, often glabrous.
Rhizomal leaves and lowermost cauline compound, pinnate, (7−)12−28 cm; petiole
(3−)6−10(−15) cm; lateral leaflets 2−4 pairs, broadly to narrowly ovate, 2−6 × 1.5−5 cm, sparsely
to moderately pubescent with trichomes to 0.5 mm, pinnately lobed or incised to coarsely dentate
or serrate, base oblique or rarely cuneate, apex acute; petiolule (0.8−)1−2.2 cm; terminal leaflet
same size or slightly larger than lateral ones; cauline leaves 3−5, progressively smaller and with
shorter petioles and petiolules upward. Racemes ebracteate, 8−15-flowered, dense corymbose,
ebracteate, slightly to strongly elongated in fruit; rachis straight; fruiting pedicels 0.9−2.5 cm,
stout, ascending to divaricate-ascending, straight. Sepals green, oblong, 4−6 × 2−2.5 mm,
caducous, glabrous; petals white, broadly obovate, (0.8−)1−1.5 cm × 4−8 mm, apex obtuse or
rounded; filaments 4−7 mm; anthers oblong, 1−1.5 mm; ovules 12−2o per ovary. Fruits linear,
(3−)4−8(−9) cm × 2−2.5(−2.9) mm, glabrous; style (2−)3−5 mm. Seeds brown, oblong, 2.5−3.2 ×
1−1.8 mm. 2n = 144.
Flowering: Oct−Feb.
Habitat: wet or swampy areas.
Elevation: 0−600 m.
Distribution: Argentina (Santa Cruz, Tierra del Fuego), Chile (Región X, XI, XII).
Specimens examined: ARGENTINA. Santa Cruz: Río Santa Cruz, Montt 197 (CONC). Dep.
Lago Argentinao, Lago Argentino, 22 Feb 1914, Hicken & Hauman s.n. (SI); Porto Ferrari, Lago
Argentino, Hicken & Hauman 375, 429 (SI). Tierra del Fuego: Dep. Ushuaia, arroyo Culebra,
10 km N Ushauaia, Boelcke et al. 15276 (B AA, BACP); Estancia Moat, 54°56′S, 66°44′W,
Moore 1693 (BAA, BAB); S Lago Fagnane, Mt. Heuhuepen, 54°34′S, 71°01′W, Moore &
Goodall 394 (BAB, K); Ushuaia, Bahía Buen Suceso, Biganzoli 467 (MO, SI); Valle Lasifashaj,
Las Cotorras, ca. 20 km ENE of Ushuaia, Santesson 416 (K, P, S); near Monte Olivia, Hunziker
8213 (CORD, GH); E Monte Olivia River of Vela de la Novia falls, Goodall & Lutken 42 (BAB,
MO); S Lago Escondido, 54°40′S, 67°53′W, Roivainen 611 (MO); Ushuaia, Monte Susana,
Roivainen 1097 (MO); Ushuaia, Pennington 307 (SI); Co. Dos Banderas, Correa & Pérez
Moreau 2058 (BAA, BAB); Valle de l’Olivaia, 14-17 Feb 1896, Alboff s.n. (SI); Parq Nac
Cañadón Toro, Correa & Reynoso 11732 (BAA); Montes Martiales, camino Laguna Azul, 9 Jan
1960, Grondona s.n. (BAA). CHILE. Ruta Austral, Paso Queulet, Schlegel 8109 (CONC);
Estancia Remolino, 54°50′S, 67°52′W, Moore 1999 (BAA, K). X. Ró Palena, Jan-Feb. 1887,
Delfuis s.n. (SGO). XI: Coihaique, El Sitio (Cerro El Cielo), Bosque Lenga, Godoy et al. 64
(CONC); Valle de río Cajón, aflueente de río Ibáñes, la parte dlta del río Ibáñez antes de su
confluencia con el río Cajón, y el valle del Estero Pérez, 46°10′S, 72°17′W,5 Dec 2002, Moreira
s.n. (SGO). XII. Punta Arenas (Sandy Point), 1878, Ortega s.n. (SGO), Dusén 107 (LIL, MO, S,
UPS), Philippi 65 (SGO), Lechler 1162 (G, P, UPS, W), Cunningham s.n. (B, GH); Punta
Arenas, Tres Brazos, [53°16′S, 70°57′W], Giorgio 37 (CONC); Río Bueno, Donat 345 (BA,
BAF, BM, CAS, F, GH, K, LIL, NY, SI); Bahia Morris, Isal Capitan Aracena, [54°15′S,
71°01′W], Pisano 3293 (BAA, CONC), Moore 2723 (BAB, K, UPS); Fiordo Peel, Río Murtillar,
50°27′S, 73°37′W, Pisano 6013 (CONC); Canal de las Montañas, 51°38′S, 73°24′W, Pisano
48
6125 (CONC); Rio Hilco, Skottsberg s.n. (S, UPS); Tierra del Fuego, Fiordioa Contralmirante
Martínez, Bahía Escandallo, Río de la Laguna, 54°13′S, 70°10′W, Pisano & Dominguez 7165
(CONC); Sector Bahía Victoria, Forestal Trillium, 54°50′S, 69°52′W, Pisano et al. 8323
(CONC). Where? : Tierra del Fuego, Rio Fontaine, 3 Jan. 1908, Skottsberg s.n. (SGO). Bahia
Morris, Isla Capitan Aracena, Pisano & Moore 3393 (BAA). Península Muñoz Gamero, Calet
Dora, Pisano & Dollenz 5879 (GH). Laguna Parrillar, Pisono 3890 (GH). Peru. Cusco, Calca,
Dist. Lares, Choquecancha, Azulcocha, 12°33′S, 72°W, Valenzuela et al. 4976 (MO).
Cardamine geraniifolia is one of the most distinctive South American species of the genus.
Despite its variation in flowers size, petiole and petiolule length, leaflet size and margin type, and
fruit length, it has not been divided into infraspecific taxa, as evidenced from the synonymy
above. It is readily distinguished from all congeners in South America by the fleshy, repent, nonscaly rhizome, pinnately compound leaves with long petiolules, 2−4 lateral pairs of serrate to
incised or lobed leaflets, and large petals to 1.5 mm long.
The above collection from Peru represents the first record of this otherwise Patagonian
species. The Peruvian plant is with immature fruits, but in floral, leaf, and receme aspects it is
indistinguishable from plants of Cardamine geraniifolia. Without mature fruits, the placement of
this collection is this species remains tentative.
Cardamine glacialis (G. Forster) de Candolle, Syst. Nat. 2: 264. 1821. Sisymbrium glaciale G.
Forster, Comm. Soc. Regiiae Göttingen 9: 32. 1789. TYPE: Argentina, Tierra del Fuego:
Ushuaia, J. R & G. Forster s.n. (holotype, BM, not seen).
Sisymbrium grandiflorum Molina, Sag. Stor. Nat. Chili, ed. 2, 292. 1810; non S. grandiflorum
Weddell, Ann. Sci. Nat., Ser. 5, 1: 290. 1864; nec S. grandiflorum Post, Pl. Postinae 1: 3.
1890. TYPE: Chile, Collector? (holotype, BOLO?). Neither type nor original publication seen.
Cardamine pratensis Linnaeus var. minor Barnéoud in Gay, Fl. Chile 1: 114. 1845. TYPE:
Chile, C. Gay s.n. (holotype, P, not seen)
Cardamine glacialis var. pumila A. Gray, A. Gray, Bot. U. S. Expl. Exped. Wilkes 1: 49. 1854.
TYPE: Tierra del Fuego, collector? (holotype, GH?, or US?).
Cardamine glacialis var. elatior A. Gray, A. Gray, Bot. U. S. Expl. Exped. Wilkes 1: 49. 1854.
TYPE: Tierra del Fuego, Orange Harbor, 1838-1842, Wilkes s.n. (holotype, GH?, or US?;
isotype, P!).
Cardamine antiscorbutica Banks & Solander ex Grisebach, Abh. Königl. Ges. Wiss. Göttingen
6: 115. 1854; C. hirsuta Linnaeus var. antiscorbutica (Banks & Solander ex Grisebach)
Reiche, Fl. Chile 1: 100. 1896 TYPE: Chile, Magallanes, Brunswick Peninsula, Sandy Point,
Lechler 1116 (lectotype, inadvertlydesignated by Steudel, GOET, not seen; isotypes, G!, NY!,
4P!, SGO!). Grisebach (1854) listed both Lechler 1116 and 1161 under the description of C.
antiscorbutica. The latter collection was taken by Stuedel as the type of C. strictula and
therefore C. antiscorbutica was lectotypified.
Cardamine strictula Steudel, Flora 39: 410. 1856. TYPE: Chile, Magellan: Sandy Point, Lechler
1161 (holotype, P!; isotypes, K!; fragment, BAA!).
Cardamine gongylodes Philippi, Linnaea 28: 664. 1856. TYPE: Chile, Chonos Island, Fr. Fonk
83 (holotype, SGO-49417!; fragments, BAA!).
Cardamine magellanica Philippi, Anal. Univ. Chile 41: 666. 1872; C. hirsuta Linnaeus var.
magellanica (Philippi) Reiche, Fl. Chile 1: 100. 1896. TYPE: Chile, Del Strecho de
Magallanes, Philippi s.n. (holotype, SGO-…). not seen, and no sheets listed by Muñoz (1960).
Cardamine litoralis Philippi, Anal. Univ. Chile 27(2): 313. 1865; C. glacialis subsp. litoralis
(Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 541. 1903. TYPE: Chile, Valdivia: boca del río
49
Hueicolla, Nov. 1864, F. Philippi 59 (holotype, SGO-63893!; fragments, BAA!). The type
information was copied from Muñoz (1960) who listed also SGO-49422 as part of the type
collection. I have not seen the latter sheet.
Cardamine palenae Philippi, Anal. Univ. Chile 81: 80. 1892. TYPE: Chile, Río Palena, Jan-Feb.
1887, Federico Delfin s.n. (holotype, SGO-71618!). Although Muñoz (1960) listed and
annotated another sheet, SGO-63892!, as part of the type collection, the sheet does not carry
Philippi’s hand writing about the collector and locality given above and, therefore, it should
not be considered as part of the type material.
Cardamine soehrensii Philippi, Anal. Univ. Chile 81: 84. 1892; C. hirsuta Linnaeus var.
soehrensii (Philippi) Reiche, Fl. Chile 1: 100. 1896; C. glacialis var. soehrensii (Philippi) O.
E. Schulz, Bot. Jarhb. Syst. 32: 543. 1903. TYPE: Chile, Andes of Prov. Santiago, horto
botanico (as H. B.), October 1890, Philippi s.n. (holotype, SGO-71636!). Muñoz (1960) listed
SGO-63892 as another specimen of the type collection, but I have not seen that.
Cardamine glacialis var. pubescens Philippi ex O. E. Schulz, Bot. Jahrb. Syst. 32: 541. 1903.
TYPE: Chile, Bridges s.n. (holotype, B!; isotype, G!).
Herbs, perennial. Rhizomes much thickened, short, forming a tuberous stem base, not scaly.
Stems (2−)7−45(−115) cm, erect, 1 to numerous from base, glabrous or sparsely to densely
pubescent throughout, slender or stout, lowermost nodes without adventitious roots. Basal leaves
and lowermost cauline compound, 2−15 cm, 3−11(−15)-foliolate, glabrous to densely pubescent;
petiole (1−)2−9(−11) cm; terminal leaflet (0.4−)0.8−3.8(−5) × 0.7−3(−4) cm, ovate to obovate or
suborbicular to subreniform, obtusely dentate to repand or entire, with petiolule 0.3−1.7(−3) cm;
lateral leaflets ovate to oblong, 0.3−2(−3.5) × 0.2−1.5(−2.5) cm, sessile or on a petiolule
0.5−4(−6) mm, entire to repand or dentate; upper cauline leaves similar to lower ones in shape
and size though sometimes smaller upwards. Racemes ebracteate; rachis straight; fruiting
pedicels 4−15(−25) mm, ascending to suberect, slender or stout, straight, often forming a straight
line with fruit. Sepals oblong to broadly ovate, (1.7−)2−3 × 0.8−1.5 mm; petals white, narrowly
obovate to oblanceolate, 4−7(−8.5) × 1.5−3 mm, gradually narrowed to base, apex obtuse;
filaments 2−4 mm; anthers ovate, 0.5−0.8 mm; ovules 20−32 per ovary. Fruits linear,
(1,6−)2−3.5(−4) cm × 1.2−2 mm, not attenuate at apex; style stout, (0.3−)0.5−1.5(−2) mm. Seeds
brown, ovate, 1−1.7 × 0.7−1 mm, wingless.
Flowering: Oct−Mar.
Habitat: moist shade under Nothofagus trees, spring seepage, swamps, costal dunes, mossy and
sandy humus over rocks.
Elevation: 0−1450 m.
Distribution: Argentina (Chubut, Santa Cruz, Tierra del Fuego), Chile (Región X, XI, XII),
Falkland Islands.
Specimens examined: ARGENTINA. Chubut: Dep. Río Senguerr, valle del lago Blanco,
Koslovsky 12493 (BAB). Santa Cruz: N Lago Rico, Glacier Natl. Park, Eyerdam et al.
24314 (DS, G, MO, SI, UC). Dep. Güer Aike, Ea. Stag River, Meseta Latorre, Co. Punta
Gruesa, 51°32′S, 72°05′W, Roig et al. 3081 (SI); Ea. Stag River, 16 km N de casco, río de los
Venados, 51°34′S, 71°58′W, TBPA 3189 (BAB, BACP, SI); Ea. Las Viscachas, Laguna
Viscachas, 50°43′S, 71°58′W, TBPA 2587 (BAB, SI); Ea. La Verdadera Argentina, Co de La
Virgen, 50°50′S, 71°14′W, TBPA 2126 (BAB, SI); canal de los Tempanos, Glaciar
Ameghino, 50°30′S, 73°0′W, Correa et al. 3101 (BAA, BAB). Dep. Lago Argentino, Lago
Argentino, Pen. Magallanes, Boelcke et al. 16431 (BAA, BAB, BACP); Porto Ferrari, Hicken
50
& Hauman 368 (SI); Estancia Cristina, N end of Lago Argentino, Ager 504 (US). Tierra Del
Fuego: Dep. to Ushuaia, Isla de los Estados, Puerto Cook, 54°45′S, 64°03′W, Nicora et al.
7349 (BAA, SI); Bahia Franklin, 54°54′S, 64°42′W, Dudley et al. 1451 (SI); Puerto Roca,
54°43′S, 64°12′W, Dudley et al. 731 (SI); Puerto Vancouver, 54°47′S, 64°04′W, Dudley et al.
1023 (A, E, MO, P, SI, UC), Dudley et al. 934 (SI); Bahia Flinders, 54°46′S, 64°32′W,
Dudley et al. 1503 (E, P); side of Lago Fagnano, Monte Heuhuepen, 54°34′S, 67°07′W,
Moore 2846 (MO); Sierra Sorondo, behind Punta Segunda, 54°47′S, 67°58′W, Moore 1959
(MO); Estancia Harberton, Monte Overo, Goodall 460 (E, NY, UC, US), Boelcke et al.15254
(BAA, BAB, BACP); Twin Islands of Estancia Haberton, Goodall 1365 (NA); Lago Fagnano,
Boelcke et al. 15204 (BAA, BAB, BACP, MO); Ushuaia, Lapataia, Boelcke et al. 15273
(BAA, BAB, BACP, MO); Lago Escondide, Cabrera 34141 (MO); Estancia La Correntina,
14 km E Fabrica Kami, Goodall 3984 (A, NA); Isla de los Estados, 54°52′S, 64°41′W,
Biganzoli 658 (SI); Mitre Peninsula of Isla Grande, 54°47-48′S, 65°15-16′W, Dudley et al.
214 (SI); Montes Martiales, camino Laguna Azul Altura, Grondona 7349 (BAA); Isal
Wollaston, Caleta Lientur, 55°44′S, 67°19′W, Pisano 4979 (GH). Dep. Islas del Atlántico
Sur, Islas Malvinas: without locality, 1839-1843, J. D. Hooker s.n. (B, G, K); Pto. Stanley,
Ulibarri et al. 238 (SI). CHILE. X: Valdivia, Corral, 10 Nov 1902, Buchtien s.n. (E, G, GH,
SI, US); Llanquihue, Isla Magdalena, Werdermann 79 (CAS, E, F, G, GH, LIL, MO, SI, UC);
Isla Grande de Chiloe, 30 km N Cucao at mouth of Rio Anay, 42°28′S, 74°15′W, Gardner
3594 (CONC, E); Chepu, Godley 343 (SGO); Pucatrihue, Sparre 3879 (SGO); Niebla,
[39°51′S, 63°24′W], Gunckel 40611 (CONC); Hueicolla, [40°09′S, 73°39′W], Hollermayer
778 (CONC, SI). XI: Portezuelo Los Mallines, enter los valles Simpson e Ibanez, Rentzell
6132 (SGO); Galeta Vidal, Behn 25 (SGO); Parque Nacional Laguna San Rafael, 46°37′S,
73°53′W, Pisano 6259 (CONC); Estero Cofré, [46°11′S, 72°47′W,] Vogel 560 (CONC);
Coihaique, Mallín, Godoy et al. 62 (CONC). XII: Isla Piazzi, Caleta Ocación, 51°44′S,
74°01′W, TBPA 1033 (SI); 15 km S Punta Arenas, Eyerdam et al. 24113 (DS, G, MO, NA,
SI, UC); 78 km NW Punta Arenas on rd to Puerto Natales, Eyerdam et al. 24144 (CAS, G,
MO, SI, UC); Estancia Maria Christina, 80 Km NE Punta Arenas, Goodall 4023 (A); Vicuña,
[54°08′S, 68°47′W], Ricardi & Matthei 231 (CONC); Punta Arenas, 24 Jan 1936, [53°10′S,
70°54′W], Behn s.n. (CONC); Cerros Canchas de Sky, [53°09′S, 70°55′W], Ricardi &
Matthei 303 (CONC); Monte Fentón, Barrientos 159 (CONC); Isla Guarelo, [50°21′S,
75°20′W], 13 Nov 1975, Moyano s.n. (CONC); Sierra Carmen silva, N Río Chico Valley,
53°30′S, 68°39′W, Moore & Goodall 219 (BAB, NA); Estancia Cameron, Moore 1105 (US);
Rio Rubens, ca. 50 Km SE Natales, Santesson 1611 (S); Isla Navarino, 2 km SW Puerto
Williams, Moore 352 (UC, US); Cerro Donoso, Sector Río de las Chinas, 50°44′S, 72°31′W,
Arroyo et al. 870308 (CONC); Sierra de los Baguales, Cerro Santa Lucía, 50°44′S, 72°20′W,
Arroyo 841121 (CONC); Parque Nacional Torres del Paine, Cerro Diente, 50°47′S, 72°57′W,
Arroyo & Squeo 850913 (CONC); Fiordo Parry, Bahia Cuevas, [54°42′S, 69°19′W], Pisano
3003, 3071 (CONC, SI); Bahia Morris, Isla Capitan Aracena, [54°15′S, 71°01′W], Pisano
30305 (CONC); Canal de las Montañas, 51°38′S, 73°24′W, Pisano 6147 (CONC); Fiordo
Peel, 50°30′S, 73°44′W, Pisano 6104 (CONC); Puerto Eden, Isal Wellington, 49°09′S,
74°33′W, Pisano 6398 (CONC); Kerber, Punto Enap, [52°45′S, 69°15′W], Pisano 2590
(BAA, CONC, SI); Río Bueno, [53°10′S, 70°00′W], Pisano 2489 (CONC); Península Roca,
51
Seno Resi, 51°51′S, 73°2′W, Pisano 2931 (SI); Fiordo Peel, Río Murtillar, 50°27′S,
73°37′W], Pisano 5897 (CONC); river outwash from Alemania Glacier, Beagle Channel,
Llano 74 (NA); Rio Santa Maria a 60 km Camino Sur, Pisano 3551 (BAA, NA); Laguna El
Parrillar, Pisano 3915 (GH, NA); Los Tres Mrros, Dolenz 183 (CONC); Lago Blanco, Texera
2 (CONC); Fiordo Silva Palma, Angostura Titus, Río Raul, Pisano 3822 (NA); Rio Caleta,
Pisano 2728 (MO, SI); Los Azules, Pisano 2737 (MO, SI); Cerro Mirador, Pisano 2769 (MO,
SI); Seno Ultima Esperanza, Puerto Bella Vista, 51°28′S, 73°16′W, Moore & Pisano 1587
(SI).
Cardamine glacialis is sometimes confused with the introduced C. hirsuta because both have
pinnate leaves and erect appressed fruits. However, C. glacialis is a perennial with tuberous stem
base, has six stamens, and blund fruits …. mm wide. By contrast, C. hirsuta is an annual with
four, rarely five or six, stamens, and fruits slener at apex and … mm wide.
Of the several infraspecific taxa recognized by Schulz (1903) in Cardamine glacialis, perhaps
subsp. litoralis might merit recognition. It differs from typical plants of the species by being
robust plants with coarser leaves, shorter styles, and wider fruits. Although most plants that
appear to belong to this variant were collected in the area Valdivia and other collections listed
above under Región X, this variation is sporadic elsewhere, and many intermediate forms
connect the two extremes of the species.
It is with a great deal of hesitation that I follow Schulz (1903) in the placement of Cardamine
soehrensii in the synonymy of C. glacialis. The plant has most aspects of the latter but the
fruiting pedicel and fruit do not form a straight line as most members of C. glacialis do.
Furthermore, the type of C. soehrensii was collected from the codillera of Santiago, an area
substantially disjunct from the distribution range of C. glacialis. It is possible that the locality
data for the type collection of C. soehrensii is erroneous, and the type was collected from a much
sourtherly area.
Schulz (1903) cited Werdermann 79 as Cardamine macrostachya (=C. vulgaris), but every
aspect of the plants of that collection are characteristics of C. glacialis. Although the species is
highly variable in the size of plant and leaf morphology, the overall aspects of the plant are
basically the same. One collection, Dudley et al. 731 (SI), is unusually tall for the species, and it
is well over one meter tall.
Of the several infraspecific taxa recognized by Schulz (1903) in Cardamine glacialis, perhaps
subsp. litoralis might merit recognition. It differs from typical plants of the species by being
robust plants with coarser leaves, shorter styles, and wider fruits. Although most plants that
appear to belong to this variant were collected in the area Valdivia and other collections listed
above under Región X, this variation is sporadic elsewhere, and many intermediate forms
connect the two extremes of the species.
Cardamine glanduligera O. Schwarz, Repert. Sp. Nov. Regni Veg. 46: 188. 1939; Dentaria
glandulosa Waldstein & Kitaibel ex Willdenow, Sp. Pl. 3: 478. 1800. TYPE: Habitat in
Hungriae sylvis subalpinis, Kitaibel s.n. (holotype, B-W 11955).
Cardamine glauca Sprengel ex de Candolle, Syst. Nat. 2 : 266. 1821. TYPE: [Italy], “in aspero
monte [Aspromonte] prope Reggio in Calabria,” L. Thomas (holotype, G-DC).
Cardamine glechomifolia H. Léveillé, Repert. Sp. Nov. Regni Veg. 11: 495. 1913. TYPE:
Korea: Quelpaert, Mokan, May 1911, Taquet 5385 (holotype, E).
52
Cardamine gouldii Al-Shehbaz, Novon 11: 289. 2001. TYPE: Bhutan, [Bumthang District],
Yatong La to Gyesta, 9,800−11,700 ft, 7 May 1938, B. J. Gould 415 (holotype, KI).
Herbs, perennial; underground parts, basal leaves, and lowermost portions of stem unknown.
Stems striate, sparsely pilose with trichomes to 0.6 mm; distal nodes 5−11 cm apart. Cauline
leaves 5-foliolate; uppermost trifoliolate; petiole 1−3 cm, narrowly winged, base not auriculate;
terminal leaflet oblong to oblong-lanceolate, 2.5−5 × 0.7−2 cm, sparsely hirsute with trichomes
to 0.6 mm, subsessile or cuneate basally into a petiolule to 5 mm, margin entire to repand,
ciliolate with trichomes to 0.1 mm, lateral veins ending in mucronulate callosities to 0.2 mm,
apex rounded to obtuse; lateral leaflets oblong, sessile, similar to and smaller than terminal
leaflet. Racemes ebracteate; lax, 5- to 10-flowered; flowering pedicels slender, recurved, 5−10
mm. Sepals drying lavender, oblong, 5−7 × 2−3 mm, sparsely pilose, base saccate; petals white,
oblong, 12−15 × 4−6 mm, apex rounded; claw ca. 1.5 mm; stamens erect; filaments of median
pairs 8−10 mm, those of lateral pair 6−7 mm; anthers oblong, ca. 1.5 mm. Fruit and seeds
unknown.
Flowering: May.
Distribution: Bhutan.
Notes: known thus far only from the type collection.
Cardamine gracilis (O. E. Schulz) T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. N. E. Forest. Inst.,
Harbin 1980(6): 27. 1980; C. multijuga Franchet var. gracilis O. E. Schulz, Repert. Spec.
Nov. Regni Veg. 17: 289. 1921. TYPE: China, Yunnan, Lichiang, near Ngu Le Keh, 2900 m,
18 Jul 1914, Camillo Schneifer 1862 (holotype, B!; isotypes, GH!, K!, US!).
Herbs, perennial, glabrous throughout, semiaquatic. Rhizomes slender, to 10 cm or longer,
0.5–3 mm in diam. Stems 1–3.5(–5) dm, erect, simple, striate angled, rooting from lower and
sometimes middle nodes. Basal leaves not known; cauline leaves 10–20 per stem, (1.5–)2.5–
7.5(–9) cm × (1.5–)4–14(–17) mm; petiole absent or rarely to 2 mm; terminal leaflet narrowly
obovate, oblong, or rarely broadly ovate, (2–)3–8(–9) × (0.5–)1.5–5(–7) mm, petiolule 0.5–2
mm, base cuneate or rarely obtuse, margin obscurely 1–3-toothed on each side, apex acute,
submucronate; lateral leaflets (6 or)7–12(–15) pairs, linear or narrowly oblong, asymmetric,
slightly smaller than terminal lobe, sessile or rarely with a petiolule to 0.5 mm, base oblique,
proximal margin 1- or 2(or 3)-toothed, distal margin entire or rarely obscurely 1- or 2-toothed,
apex acute; proximal pair of lateral leaflets auriclelike, attached at or just above node, often
giving appearance of amplexicaul leaf base. Racemes ebracteate, 10–20-flowered; fruiting
pedicels divaricate, 0.8–2.2 cm, slender. Sepals oblong or ovate, 3–4 × 1.2–2 mm, spreading,
margin membranous, base of lateral pair subsaccate; petals lavender with darker veins, spatulate,
6–8 × 2–3 mm, spreading, apex rounded; filaments spreading, dilated at base, subequal, median
pairs 3.5–5 mm, lateral pair 3–4.5 mm; anthers narrowly oblong, 1.3–1.6 mm; ovules 10–14 per
ovary. Fruit linear, 2–4 cm × ca. 1.5 mm; gynophore ca. 0.5 mm; valves smooth, glabrous; style
4–6 mm. Seeds brown, oblong, 1.7–2 × ca. 1.3 mm, winged all around; wing ca. 0.4 mm wide.
Flowering: Jun–Aug.
Habitat: marshlands, lake shores, margins of ponds, ditches, open pastures.
Elevation: 2400–3300 m.
Distribution: China (Yunnan).
Cardamine graeca Linnaeus, Sp. Pl. 2: 655. 1753. TYPE: Herb. A. Van Royen, sheet 901.22060. (lectotype designated by Marhold (1996: 118), L).
53
Cardamine granulifera (Franchet) Diels, Notes Roy. Bot. Gard. Edinburgh 5: 204. 1912. C.
tenuifolia (Ledebour) Turczaninow var. granulifera Franchet, Bull. Soc. Bot. France 33: 399.
1886; Loxostemon granulifer (Franchet) O. E. Schulz, ……. TYPE: China, Yunnan, Mt.
Tsang-chan, above Tali, 2 Jun 1883, Delavay 281 (holotype, P!).
Herbs, perennial, glabrous or sparsely pilose. Rhizomes much elongated, slender; bulbils
few, spaced, rarely clustered. Stems 6–20(–30) cm, simple, erect, straight, slender, sparsely
pilose along proximal half, or glabrous. Rhizomal leaves simple or trifoliolate; petiole 0.7–4 cm;
leaf blade or terminal leaflet suborbicular, broadly ovate, or cordate, 3–12 mm in diam., base
cordate or rounded, margin entire or obscurely lobed; cauline leaves 2–4, trifid or pinnatisect and
with 2(or 3) lateral lobes on each side of midvein, lobes not decurrent; petiole 1–15(–28) mm,
slender, wingless, glabrous, not auriculate at base; terminal lobe filiform to narrowly linear, 0.9–
4.5 cm × 0.5–1.5(–2) mm, base attenuate, margin entire, scabrous with trichomes to 0.3 mm,
apex acute. Racemes ebracteate, 3–10-flowered; flowering pedicels ascending, to 1.2 cm,
straight, glabrous. Sepals ovate, 2–1.5 × 1–1.5 mm, glabrous, margin and apex membranous,
lateral pair subsaccate; petals white or lavender, broadly obovate or spatulate, 5–7 × 2–3 mm,
apex roundedl; median filament pairs 3–4, slightly flattened; lateral pair 1.5–2.5 mm; anthers
oblong, 0.6–0.8 mm. Fruit and seeds not seen.
Flowering: May–Jul.
Habitat: Moist shady forests, meadows.
Elevation: 2800–3800 m.
Distribution: China (Yunnan).
Cardamine granulose Allioni
Cardamine griffithii J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5: 146. 1861. TYPE:
Bhutan, Lamnoo, Griffith s.n. (holotype, K!).
Cardamine griffithii var. pentaloba W. T. Wang, Acta Bot. Yunnan. 9: 16. 1987. TYPE: China.
Yunnan: Lanping, 2600−3100 m, 28 Jun 1981, Exped. Hengduanshan 1171 (holotype, PE!).
Herbs, perennial, glabrous throughout except for leaflet margin. Rhizomes creeping, without
stolons. Stems (2–)3–10(–11.5) dm, erect, simple or branched above, striate angled, (9–)12–28(–
37)-leaved. Leaves sessile, cauline; lower and middle ones (1–)2–9(–11) × (0.7–)1–3.5(–4.5) cm;
terminal leaflet orbicular, broadly ovate, or obovate, (0.5–)1–3(–3.5) × (0.3–)0.6–1.9(–2.5) cm,
with a petiolule 2–10(–15) mm, base subcordate, rounded, or rarely cuneate, margin repand or
entire and sparsely ciliate, apex obtuse or rounded; lateral leaflets 2–4(or 5) pairs, slightly to
distinctly smaller than terminal one, base obtuse or rarely slightly oblique, margin entire or
repand and sparsely ciliate, apex obtuse or rounded; proximal pair of lateral leaflets auriclelike,
attached at or just above node, often giving appearance of amplexicaul leaf base; uppermost
leaves smaller. Racemes ebracteate; fruiting pedicels ascending or divaricate, (0.4–)0.7–1.5 cm,
slender, straight. Sepals ovate or oblong, 2.5–3 × 1.5–2 mm, erect; petals purple or lavender,
spatulate or obovate, 6–9 × (2.5–)3–5 mm, not clawed, apex rounded or subemarginate; filaments
erect, median pairs 3.5–4.5 mm, lateral pair 2.5–3 mm; anthers oblong, 0.9–1.1 mm; ovules 10–
22 per ovary. Fruit linear, (1.5–)2–4 cm × 0.9–1.2 mm; gynophore 0.5–1 mm; valves smooth,
glabrous; style 0.5–1(–2) mm; stigma 2-lobed, distinctly broader than style. Seeds brown,
oblong, 1.4–1.7 × 0.8–1.1 mm, wingless.
Flowering: May–Aug.
54
Habitat: mountain slopes, valleys, streamsides, pastures, marshy places, moist forest floor, shady
rocky areas.
Elevation: 2400–4500 m.
Distribution: Bhutan, China (Sichuan, Xizang, Yunnan), India (Sikkim), Nepal.
Cardamine gunnii Hewson, Fl. Austral. 8: 390. 1982; Cardamine heterophylla W. J. Hooker,
Comp. Bot. Mag. 1: 273. 1835; not Host, Syn. P. Austral. 366. 1797; not Lapeyrouse, Hist.
Abr. Pl. Pyr. 377. 1813; not Bory, Ann. Sci. Gen. Phys. 3: 6. 1820; Cardamine hirsuta var.
heterophylla (W. J. Hooker) J. D. Hooker, Fl. Tasmaniae 1: 20. 1855. TYPE: Australia,
Tasmania, R. C. Gunn 446 (lectotype designated by Hewson (1982: 390), K).
Herbs perennial, glabrous or rarely pubescent, often scapose. Rhizomes present. Stems to 25
cm, leafless or rarely leafy, erect. Basal leaves rosulate, pinnately compound with 1 to several
pairs of leaflets, pinnatisect, dentate, or entire, spatulate; cauline leaves (when present) petiolate,
not auriculate at base. Raceme ebracteate, corymbose; fruiting pedicels erect to spreading, 5–17
mm, often minutely papillose. Sepals 2–2.5 mm, purplish; petals 4–6 mm, white; stamens 6;
ovules –. Fruits linear, 1.5–3 cm × 0.7–1 mm, erect, often overtopping raceme; style < 1 mm.
Seeds ovoid to oblong, flattened, 1–1.5 mm, narrowly winged.
Habitat: moist areas in alpine regions down to coastal places.
Distribution: Australia (New South Wales, South Australia, Tasmania, Victoria).
Cardamine heptaphylla (Villars) O. E. Schulz, Bot. Jahrb. Syst. 32: 371. 1903; Dentaria
heptaphylla Villars, Hist. Pl. Dauphiné 1: 281. 1786. TYPE: Dentaria eptaphyllos [Villars]
(lectotype designated by Marhold (2001: 47), GR, no. MNHGr. 1837.27769).
Cardamine hirsuta Linnaeus, Sp. Pl. 2: 655. 1753. TYPE: Herb. Linn. 835.13 (lectotype,
designated by Fawcett & Rendle (Fl. Jamaica 1914: 239), LINN).
Cardamine hirsuta var. formosana Hayata, Mat. Fl. Formos. 30. 1911.
Cardamine hirsuta var. rotundiloba Hayata, Mat. Fl. Formos. 30. 1911.
Herbs, annual, sparsely hirsute at least along petioles of basal leaves, often glabrous above.
Rhizomes absent; stolons absent. Stems (0.3–)1–3.5(–4.5) dm, erect, ascending, or decumbent, 1
to several from base, simple or branched above, not flexuous. Basal leaves rosulate, persistent by
anthesis, (5–)8–15(–22)-foliolate, (2–)3.5–15(–17) cm; petiole ciliate, 0.5–5 cm; terminal leaflet
reniform or orbicular, 0.4–2 × 0.6–3 cm, margin entire, repand, dentate, or 3–5-lobed, petiolule
2–10 mm; lateral leaflets oblong, ovate, obovate, or orbicular, smaller than terminal lobe, entire,
repand, crenate, or 3-lobed; cauline leaves 1–4(–6), compound as basal leaves, shortly petiolate,
including petiole (0.5–)1.2–5.5(–7) cm; petiole base not auriculate. Racemes ebracteate; fruiting
pedicels erect to ascending, (2–)3–10(–14) mm. Sepals oblong, 1.5–2.5 × 0.3–0.7 mm, base of
lateral pair not sacate; petals white, spatulate, 2.5–4.5(–5) × 0.5–1.1 mm, sometimes absent;
stamens 4 and lateral pair often absent, rarely 5 or 6; filaments 1.8–3 mm; anthers ovate, 0.3–0.5
mm. Fruit linear, (0.9–)1.5–2.5(–2.8) cm × (0.8–)1–1.4 mm, often appressed; valves glabrous,
torulose; style 0.1–0.6(–1) mm; ovules and seeds 14–40 per fruit. Seeds light brown, oblong or
subquadrate, 0.9–1.3(–1.5) × 0.6–0.9(–1.1) mm, narrowly margined. 2n = 16.
Flowering: Feb–Jul.
Habitat: roadsides, clearings, disturbed sites, slopes, cedar glades, mixed woods, meadows,
fields, waste ground, damp places, grassy areas, paramo.
Elevation: 0–3600 m.
Distribution: native of Europe (all countries) and perhaps W Asia (Armenia, Azerbaijan, Cyprus,
Georgia, Iran, Iraq, Israel, Jordan, Lebanon, Saudi Arabia, Syria, Turkmenistan, Turkey);
55
naturalized in S Africa, elsewhere in Asia [China (Yunnan), India, Indonesia, Japan, Korea,
Laos, Malaysia, Papua New Guinea, Pakistan, Philippines, Sri Lanka, Thailand, Vietnam],
Australia, North America [Canada (British Columbia, Ontario), United States (Alabama,
Arkansas, California, Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois,
Kentucky, Louisiana, Maryland, Massachusetts, Mississippi, Missouri, New Jersey, New
York, North Carolina, Ohio, Oklahoma, Oregon, Pennsylvania, South Carolina, Tennessee,
Texas, Utah, Virgina, West Virginia, Washington)], and South America [Argentina (Buenos
Aires, Chubut, Entre Rios, Salta, Neuquén, Santa Cruz, Tucumán), Brazil (Rio Grande do
Sul), Chile (Región V, Santiago, VI, VII, VII, VIII, IX, X, XII), Columbia (Antioquia),
Ecuador (Cotopaxi, León, Napo, Pichincha), Uruguay (Montevidio), Venezuela (Aragua)].
Representative specimens: ARGENTINA. Buenos Aires: Tigre, Burkart 3886 (SI); La Plata,
Burkart 749 (SI); La Plata, Vivero del Bosque Lugare Sombreado, Cabrera 4527 (F, GH, NY,
SI); Gral Madariaga, Vill Gesell, Boelcke 8725 (K); San Miguel, 15 Sep 1960, Burkart s.n. (SI);
Delta del Paraná, Burkart 4943 (SI); Tandil, Boelcke 8654 (BAA). Chubut: Dep. Futaleufú,
Esquel, Burkart 19090 (SI); Trevelín, Soriano 4440 (BAA). Dep. Futaleufú: ruta 258, 24 km S
lago Rosario, Correa et al. 9003 (BAB). Entre Rios: Concepcion del Uruguay, Burkart 23587
(SI). Salta: La Caldera, rte 9, 2 km N Ojoa de Agua, 5 km S of Abra de Santa Laura, Novara
8037 (G, S); Caldera, Abra de los Sauces, Cabrera 34137 (SI). Neuquén: Dep. Los Lagos, Parq.
Nac. Nahuel Huapi, Puerto Manzano, Boelcke & Correa 11735 (BAA); pla. Qutrihué, Boelcke &
Correa 6294 (BAA, BAB). Río Negro: Dep. Bariloche, co. Bella Vista, Barba 2214 (LIL).
Santa Cruz: N end Lago Argentino, Estancia Cristina, Ager 522 (US). Tucumán: Tafi, Cerro
San Savier, Villo 50 (BAB, F, GH); Quebrada de Lules, Venturi 928 (LIL, SI), Parodi 5230
(BAA); Cuest del Clavillo, 21 July 1981, Martínez s.n. (BACP). BRAZIL. Rio Grande do Sul:
Mun. de Porto Alegre, Bueno 3665 (F). CHILE. V: Cerro Huinco, Dan Francisco de Limache,
Looser 144 (GH); Limache, Garaventa 940 (GH, US); Cajón de San Pedro, “Mal Paso”,
[32°57′S, 71°11′W], Garaventa 4598 (BACP, CONC, SI); Cerro de la Campana, [32°55′S,
71°08′W], Garaventa 2869 (BAA, BACP, CONC, SI); Limache, Estero de Maitenes, 32°59′S,
71°15′W], Garaventa 3381 (CONC); Quebrada del Almendro, 33°0′S, 71°16′W], 3 Nov. 1916,
Behn s.n. (CONC); Palmas de Alvarado, [33°3′S, 71°06′W], Garaventa 4518 (BACP, CONC,
SI); Estero de Limache, Garaventa 2397 (BAA, BACP, SI); La Huinca, Garaventa 1715 (SI);
Estero de Maitenes, Garaventa 400 (BAA, BACP, SI); Masatierra, San Juan Bautista, 8 July
2002, Cuevas s.n. (CONC). SANTIAGO: Santiago, Claude-Joseph 1361 (US); Peñalolén,
[33°28′S, 70°32′W], Gunckel 23311 (CONC), Gunckel 25910 (CONC), Gunckel 4478 (CONC);
Marga-Marga valley, Zöllner 9363 (NA). VI: Baños de Caucuenes, [34°15′S, 70°34′W],
Gunckel 39056 (CONC); Colchagua, San Fernando, [34°32′S, 71°1′W], Montero 3601 (CONC).
VII: Talca, Claude-Joseph 2048 (US); Curicó, Cerro Condell, Barros 6442 (US); Rauquén,
34°57′S, 71°12′W, Reyes 3 (CONC). VIII: Chillán, [36°36′S, 72°05′W], Matthei 27149, 27150
(CONC); Bureo, Chillán, [36°34′S, 71°46′W], Barros 2787 (CONC, SI); Curapalihue, 36°49′S,
72°50′W, Olea 16 (CONC); Concepción, Cerro Carocol, 36°50′S, 73°02′W, Arriagada &
Ugarte 3 (CONC); Concepción, [36°49′S, 73°03′W], Barros 1966 (CONC); Parquie Hualpén,
[36°47′S, 73°10′W], Carrasco 149 (CONC). IX: Temuco, [38°44′S, 72°33′W], Montero 2344
(CONC, SI), Gunckel 16549 (CONC); Cerro Ñielol, [38°43′S, 72°35′W], Montero 5212
(CONC); Nuevo Imperial, above Cholchol, [38°36′S, 72°51′W], Montero 11878 (CONC); Licán
Ray, [39°29′S, 72°09′W], Montero 12566 (CONC). X: Corral, Gunckel 2341 (GH); Trumao, La
Unión, Hollermayer 637b (CONC). XII: Parque Nacional El Paine, 10 km from Sarmiento, Nov.
1988, Dombrowicz s.n. (G). COLOMBIA. Antioquia: Medellín, Corregimiento Santa Helena,
56
Vereda La Quiebra, 21 km SE Medellín, 6°14′N, 75°30′W, Callejas 9567 (NY). Caldas. road
from Fresno to Manizales, Gentry et al. 34746 (MO). ECUADOR. Cotopaxi: Cantolon
Sigchos, Campo Alegra, ca. 20 km NW of Sigchos, 0°34′S, 78°47′W, Ramos et al. 5905 (MO);
Parque Nac. de Cotopaxi, NW Limpio Pungu to foot of Volcán Rumiñahui, 0°38′S, 78°28′W,
Laegaard 101449 (AAU, MO). León: Cotopaxi railway station, Asplund 6511 (CAS, G, K, NY,
S). Napo: Cantón Quijos, 4 km N Papllacta, 0°19′S, 78°8′W, Yánez & Maza 930 (MO); QuitoLago Agrio rd, W Laguna Papalacta, Balslev & Steere 4533 (NY). Pichincha: Quito, Loma
Murillo, 1 km fro rio Alambi, 0°2′44″S, 78°37′52″W, Delgado et al. 105 (MO); Parroquia
Calacalí, 0°5′N, 78°30′W, Cerón 1813 (MO); Salvador, below San Juan, Asplund 17153 (G,
NY); below San Juan towards Chiriboga, Asplund 16237 (G, K, NY); between km 37 and 50
along Río Saloya, Steyermark 52496 (K, NY). URUGUAY. Montevideo: Miguelete, Herter
80831 (F, G, GH, LIL, MO, NY, S, SI, UC); Prado, Rosengurt 1955 (GH) VENEZUELA.
Aragua: Monumento Natural Pico Codazzi, 4 km WSW Colonia Tovar en la carreteras Los
Anaucos, 10°235′N, 67°19′W, Meier et al. 6489 (MO); 7 km SE Colonia Tovar, 10°21′N,
67°14′W, Steyermark & Liesner 121852 (MO, NY).
A highly variable species, especially with respect to shape, size, margins, and number lateral
leaf lobes, plant size, density of indumentum, and flower morphology (e.g., presence vs. absence
of petals, stamen number). The species has been divided into more than 15 subspecies, varieties,
and forms. However, all of these taxa are based on characters that can be highly variable even
within the same population. Only one synonym pertinent to China is cited above. For additional
synonymy, the reader should consult Schulz (Bot. Jahrb. Syst. 32: 464–473. 1903).
Cardamine hispidula Philippi, Anal. Univ. Chile 81: 79. 1892; C. alsophila Philippi var.
hispidula (Philippi) Reiche, Fl. Chile 1: 99. 1896. TYPE: Chile, Andes of Prov. Ñuble, October
1878, Fed. Puga s.n. (holotype, not seen).
Herbs, perennial. Rhizome globose-tuberous, stolon filiform, tuberous at apex. Stems 2–15
cm, erect, simple or branched at base, 8–15-leaved, flexuous, slender, subanglular, sparsely
hirsute to glabrescent. Leaves minute, those of the rhizome rosulate, 1.2–3 cm, petiolate, 3foliolate; terminal leaflet orbiculate, base subcoredate, obscurely crenate or 6-angled, long
petiolulate, 3.5–8 × 2.5–8 mm, lateral leaflets ovate, entire, short petiolulate; stem leaves 0.8–2.5
cm, lower ones 3–5-foliolate; terminal leaflet ovate, strongly 1–2 crante, short petiolualte, 2.5–8
× 1.5–6 mm; lateral leaflets similar but sessile; uppermost leafes in racemes, short petiolate, 3foliolate, leaflets 1-toothed; uppermost subsessile, simple, lanceolate, entire or 1-toothed; hirsute
to long ciliate. Racemes ebracteate, 10-flowered, flowers small, ca. 3 mm. Ovary 20–28 ovuled.
Fruit 1.8 cm × 0.7 mm.
Distribution: Chile (Región X?).
Note: known only from the type collection.
Cardamine holmgrenii Al-Shehbaz, Harvard Pap. Bot. 11: 275. 2007. TYPE: United States,
Oregon, Baker County: Blue Mountains, boggy slopes 3 miles below (north) Anthony Lake,
Elkhorn Ridge, 9 Aug 1946, Basset Maguire & Arthur H. Holmgren 26941 (holotype, NY!;
isotype, UTC).
Herbs, perennial, glabrous throughout. Rhizomes slender, 0.5–1 mm in diam.; stolons absent.
Stems 1.5–2 dm, erect to ascending. Leaves all cauline, 3 or 4, compound, 3- or 5-foliolate;
petiole 0.5–1.7 cm, base not auriculate; terminal leaflet obovate to lanceolate, becoming linear in
uppermost leaves, 1.3–2.3 cm × 2–8 mm, base cuneate, margin entire subapically 1- or 2-toothed,
petiolule 3–5 mm; lateral leaflets linear to linear-lanceolate, 0.9–1.5 cm × 1–3 mm, entire.
57
Racemes ebracteate; fruiting pedicels ascending to suberect, 5–14 mm. Sepals oblong, 1.2–1.5 ×
0.6–0.8 mm, erect, base of lateral pair not saccate; petals white, oblanceolate, 2–2.5 × 0.8–1 mm,
not clawed, apex rounded; median filament pairs 1.5–1.8 mm, lateral pair 1–1.2 mm; anthers
ovate, 0.2–0.3 mm. Fruit linear, 1–2 cm × ca. 1 mm; valves glabrous; style 0.5–0.7 mm; ovules
and seeds 16–24 per fruit. Seeds brown, oblong, 1.2–1.5 × 0.8–1 mm.
Flowering: Jul–Aug.
Habitat: boggy slopes.
Distribution: United States (E Oregon/Baker Co.).
Notes: known only form the type gathering.
Cardamine hydrocotyloides W. T. Wang, Acta Bot. Yunnan. 9: 8. 1987. China, Yunnan,
Deqin, Benzilan, Mons Yongzhongshan, 6 Jul 1981, 3200–3400 m, Qinghai-Xizang Team
2175 (holotype, PE!; isotypes, KUN!).
Herbs, perennial. Rhizomes short, thickened and somewhat fleshy at stem base, with several
stolons bearing simple leaves. Stems 10–26 cm, erect, subglabrous or puberulent with often
curved trichomes, simple from rhizomal branches, not flexuous. Rhizomal leaves and lowermost
cauline simple; petiole 1.5–6 cm, puberulent or glabrous, not auriculate at base; leaf blade
reniform or suborbicular, 0.5–2.5 × 0.8–3.5 cm, papery, glabrous or sparsely pilose adaxially,
base cordate, margin repand-crenate or subentire; stolon leaves simple, similar to rhizomal leaves
but smaller and with shorter petioles; uppermost cauline leaves usually 3-foliolate, rarely simple;
petiole 0.3–2 cm; terminal leaflet suborbicular, reniform, or ovate-suborbicular, 0.6–2 × 0.8–3
cm, with a petiolule 0.5–7 mm, base cordate, rounded, or subtruncate, margin entire or repand,
rarely slightly lobed; lateral leaflets similar to terminal one, petiolulate. Racemes ebracteate;
fruiting pedicels 0.9–2 cm, divaricate or ascending, slender, glabrous or puberulent. Sepals
ovate, 2–2.5 × 1–1.5 mm, glabrous, not saccate; petals white, obovate, 5–6 × 3–4 mm, apex
rounded; median filament pairs 3–3.5 mm, slender, toothless; lateral pair 2–2.5 mm; anthers
oblong, 0.8–1 mm; ovules 10–14 per ovary. Fruit linear, 1–2.8 cm × 1–1.3 mm; valves glabrous,
smooth; style 1–3 mm. Seeds brown, oblong, ca. 1.3 × 0.8 mm, wingless.
Flowering: Jul−Aug.
Habitat: forests, along ditches and trails.
Elevation: 3200–3400 m.
Distribution: China (Sichuan, Yunnan).
Cardamine hygrophila T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. N. E. Forest. Inst., Harbin
1980(6): 26. 1980. TYPE: China, Sichuan, Nanchuan Xian, Jinfushan, 1,650, 14 Apr 1957,
Yong Jihua 90292 (holotype, IBSC; isotype, PE!).
Herbs, perennial. Rhizomes compact, thick, not stoloniferous. Stems (7–)10–25(–35) cm,
erect, terete, pilose at base, glabrous from the middle upward, often more than 1 from base. Basal
leaves rosulate, (3–)5–9(–11) cm; petiole (0.7–)1.5–3.5(–4.5) cm; terminal leaflet orbicular or
reniform, (0.5–)1–3 × (0.5–)1.5–3.5 cm, with a petiolule 1–2 cm, base cordate or subtruncate,
margin entire, repand, or obscurely coarsely crenate, apex rounded; lateral leaflets 2 or 3(–5)
pairs, much smaller than terminal one; cauline leaves 2–5(–7), glabrous; middle ones 1.5–6(–8)
cm; terminal leaflet suborbicular or broadly ovate, rarely oblong or sublanceolate, (0.5–)1.2–3(–
3.5) × (0.5–)1–2.5(–3) cm, with a petiolule 5–12(–20) mm, base rounded or subcuneate, margin
obscurely to strongly angled, rarely entire, often mucronulate at angles, apex obtuse to subacute;
lateral leaflets (1 or)2 or 3 pairs, smaller and somewhat similar to terminal ones; proximal pair of
lateral leaflets auriclelike, attached at or just above node, often giving appearance of amplexicaul
58
leaf base. Racemes ebracteate; fruiting pedicels divaricate or ascending, (0.7–)1–2.2(–3) cm,
slender, glabrous, straight. Sepals ovate or oblong, 2.5–3 × 2–2.5 mm; petals white, obovate, 6–9
× 3–4 mm, not clawed, apex rounded; filaments subequal in length, somewhat spreading, 4.5–6
mm; anthers narrowly oblong, 1–1.3 mm; ovules 12–22 per ovary. Fruit linear, 1.5–4 cm × 1–
1.5 mm; gynophore to 1 mm; valves smooth, glabrous; style slender, 2–4 mm; stigma entire,
narrower than style. Seeds brown, oblong, 1.3–1.8 × 0.7–1 mm.
Flowering: Apr–Jun.
Habitat: valleys, streamsides.
Elevation: 1400–2200 m.
Distribution: China (Guangxi, Guizhou, Hubei, Hunan, Sichuan).
Cardamine iliciana
Cardamine impatiens Linnaeus, Sp. Pl. 2: 655. 1753. TYPE: Herb. Linn. 835.9 (lectotype
designated by Jafri (1973: 169), LINN).
Cardamine basisagittata W. T. Wang, Acta Bot. Yunnan. 9: 10. 1987. TYPE: China, Sichuan,
Kangding, Dajiangwan, 2700 m, 27 May 1974, Exped. Bot. Sichuan 6079 (holotype, PE!).
C. dasycarpa Marschall von Bieberstein, Fl. Taur. Cauc. 3: 437. 1819; C. impatiens var.
dasycarpa (Marschall von Bieberstein) T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. North-East.
Forest. Inst., Harbin 1980(6): 21. 1980. TYPE:
Cardamine glaphyropoda O. E. Schulz, Acta Hort. Gothob. 1: 159. 1924. LECTOTYPE (here
designated): China, Sichuan, Chunche, W of Sungpan, 32°39′N, 102°58′E, 1 August 1922,
Harry Smith 4121 (hololectotype, UPS!; isolectotypes, B!, W!).
C. glaphyropoda var. crenata T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. North-East. Forest. Inst.,
Harbin 1980(6): 21. 1980. TYPE: China, Sichuan, Markam Xian, 2520 m, 29 May 1957, Li
Xing 70889 (holotype, SZ; isotype, NAS!).
Cardamine impatiens Linnaeus var. angustifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 459. 1903.
Cardamine impatiens Linnaeus var. dasycarpa (M. Bieb.) T. Y. Cheo & R. C. Fang, Bull. Bot.
Lab. North-East. Forest. Inst., Harbin 6: 21. 1980. TYPE:
Cardamine impatiens Linnaeus subsp. elongata O. E. Schulz, Bot. Jahrb. Syst. 32: 459. 1903.
Cardamine impatiens Linnaeus var. eriocarpa de Candolle, Syst. Nat. 2: 262. 1821.
Cardamine impatiens Linnaeus var. fumaria H. Léveillé, Repert. Sp. Nov. Regni Veg. 12: 100.
1913.
Cardamine impatiens Linnaeus var. microphylla O. E. Schulz, Bot. Jahrb. Syst. 32: 460. 1903.
Cardamine impatiens Linnaeus var. obtusifolia Knaf, Flora 29: 294. 1846.
Cardamine impatiens Linnaeus var. pilosa O. E. Schulz, Bot. Jahrb. Syst. 32: 459. 1903.
Cardamine impatiens Linnaeus var. tenuissima Honda, Bot. Mag. (Tokyo) 54: 1. 1940.
Cardamine nakaiana H. Léveillé, Repert. Sp. Nov. Regni Veg. 10: 350. 1912. TYPE: Korea,
Quelpaert, Tpjengen, 11 Jun 1910, Taquet 4116 (holotype, E!).
Cardamine senanensis Franchet & Savatier, Enum. P. Jap. 2: 280. 1879. Japan, Prov. Senano,
Savatier 2803 (holotype, P!).
Herbs, biennial or rarely annual, glabrous or rarely sparsely pubescent near base. Rhizomes
absent; stolons absent. Stems (1.2–)2–6.5(–9) dm, erect, simple at base, usually branched, above,
angled, sometimes flexuous. Basal leaves rosulate, often withered by flowering, similar to
cauline leaves but with fewer later leaflets; cauline leaves to 9–24, (9–)13–25-foliolate, including
petiole (3–)4–20(–23) cm; petiole 2–6 cm, auriculate, the auricles to 10 × 2.2 mm; terminal lobe
orbicular, obovate, ovate, or lanceolate, 1–4(–5) × 0.5–1.7 cm, with a petiolule to 5 mm, entire or
obscurely to strongly 3–5(–9)-toothed or -lobed; lateral lobes similar to terminal one, often
59
smaller. Racemes ebracteate; fruiting pedicels divaricate or ascending, 3.5–12(–15) mm. Sepals
oblong, 1.2–2(–2.5) × 0.7–1(–1.2) mm, base of lateral pair not saccate; petals white,
oblanceolate, 1.5–4(–5) × 0.6–1.2 mm, rarely absent; stamens 6; filaments 2–3(–4) mm; anthers
ovate, 0.3–0.5 mm. Fruit linear, (1–)1.6–3(–3.5) cm × 0.9–1.5 mm; valves glabrous or rarely
pilose, torulose; style 0.6–1.6(–2) mm; ovules and seeds 10–30 per fruit. Seeds brown, oblong,
1.1–1.5 × 0.8–1 mm, compressed, sometimes apically narrowly winged. 2n = 16.
Flowering: Feb–Jul.
Habitat: streamsides, slopes, roadsides, fields, disturbed areas.
Elevation: 0–4000 m.
Distribution: native to Europe (all countries except Iceland and Portugal) and Asia [Afghanistan,
Bhutan, China (Anhui, Fujian, Gansu, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Jilin,
Liaoning, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan),
India, Japan, Kashmir, Kazakhstan, Korea, Kyrgyzstan, Nepal, Pakistan, Russia, Taiwan,
Tajikistan, Uzbekistan]; naturalized in South Africa, North America [Canada (Ontario), United
States (Connecticut, Kentucky, Michigan, Minnesota, New Hampshire, Ohio, Pennsylvania,
Virginia, West Virginia)].
Cardamine jamesonii Hooker, London J. Bot. 6: 293. 1847. TYPE: Ecuador, Cordillera of
Pilaro, 15,000 ft, Apr 1834, W. Jameson 88 (lectotype, here designated, K!).
Cardamine punicea Turczaninow, Bull. Soc. Imp. Naturalistes Mocou 27(2, 4): 295. 1854
(1855); C. johnstonii Oliver prol. punicea (Turczaninow) O. E. Schulz, Bot. Jahrb. Syst. 32:
420. 1903. TYPE: Venezuela, Merida, Culata, 9−10,000 ft, Apr 1847, Funck & Schlim 1542
(holotype, KW; isotypes, G!, P!).
Cardamine nevadensis Turczaninow, Bull. Soc. Imp. Naturalistes Mocou 27(2, 4): 295. 1854; C.
jamesonii Hooker var. nevadensis (Turczaninow) O. E. Schulz, Bot. Jahrb. Syst. 32: 422.
1903. TYPE: Venezuela, Merida, Sierra Nevada, 9,000 ft, Jun 1847, Funck & Schlim 1554
(holotype, KW?; isotypes, G!, 2P!).
Cardamine pulchra Linden & Planchon, Trois. Voy. Linden, Bot., Pl. Columb. 1: 12. 1863.
TYPE: Venezuela, Merida, Culata, 9−10,000 ft, Apr 1847, Funck & Schlim 1542 (holotype,
P!; isotypes, G!, KW).
Cardamine tolimensis Planchon & Linden in Triana & Linden, Ann. Sci. Nat. Bot. ser. 4, 17: 59.
1862. TYPE: Colombia, Pic de Tolima, près de Boqueron, 3300−3900 m, Jun 1843, Linden
924 (922 in print) (lectotype, here designated, P!).
Cardamine jamesonii var. goudotii Planchon & Linden in Triana & Linden, Ann. Sci. Nat. Bot.
ser. 4, 17: 59. 1862. TYPE: Colombia, Mariquita: Toluma, Cuchilla de la Divisadera, 1844, J.
Goudot (lectotype, here designated, P!).
Cardamine jamesonii prol. pulcherrima O. E. Schulz, Bot. Jahrb. Syst. 32: 422. 1903. TYPE:
Colombia, Cauca: Páramo de Guanacas, 2800−3300 m, F. C. Lehmann 5605 (holotype, B!;
isotypes, G!, K!).
Cardamine johnstonii var. superba O. E. Schulz, Bot. Jahrb. Syst. 32: 421. 1903. TYPE:
Colombia, Cauca: Páramo de Moras, 3300 m, F. C. Lehmann 2669 (holotype, G-BOIS).
Cardamine rhizomata Rollins, J. Arnold Arbor. 21: 392. 1940. TYPE: Ecuador, Carchi, Nudo de
Boliche Voladero, 10 Jun 1939, C. W. Penland & R. H. Summers 870 (holotype, GH!; isotype,
COCO).
Herbs, perennial. Rhizomes slender, not scaly, with long internodes and nontuberous nodes.
Stems (1.5−)2.8−6.5(−8) dm, erect or decumbent, simple or branched above, glabrous or rarely
pubescent. Cauline leaves 6−20; lower and middle leaves (4−)7−14(−18) cm, 7−11(−13)-
60
foliolate, rarely basal ones 5-foliolate; petiole (1.5−)2.5−8(−10) cm, sometimes longer on
rhizomal leaves; lateral leaflet (0.5−)1.5−3(−4.5) × (0.3−)1−2.2(−3) cm, broadly ovate to oblongovate or oblong-lanceolate, glabrous or sparsely to densely pubescent, base strongly oblique and
subcordate to cuneate, margin crenate, incised-crenate, or rarely serrate, ciliolate with trichomes
to 0.5 mm, teeth mucronate, apex acute; petiolule 0.2−1(−2) cm; terminal leaflet about same size
as lateral; uppermost leaves gradually smaller, less divided, and often narrower. Racemes
bracteate basally or very rarely only lowermost few flowers bracteate; bracts simple or
compound, gradually reduced in size upward; fruiting pedicels (1−)1.5−3 cm, divaricateascending, straight, stout. Sepals dark purple, broadly oblong, 3−5 × 2−2.5 mm, glabrous; petals
purple, violet, red, or rarely pale lavender, broadly obovate, (8−)10−14 × 5−8 mm, base broadly
cuneate, apex rounded to emarginated; filaments 4−7 mm; anthers oblong, 1−1.5 mm; ovules
16−28 per ovary. Fruits 3−5.5(−7) cm × 1.5−2 mm; style stout, 3−8(−10) mm. Seeds
oblong,1.5−3 × 1−1.2 mm.
Flowering: Sep−Jul.
Habitat: wet trails in forests, páramo, moist banks of quebrada, heath and chapparal vegetationm
mixed shrubby paramo, dwarf woodland.
Elevation: 2500−3850 m.
Distribtution: Colombia (Cauca, Mariquita, Nariño, Putumayo, Quindio, Tolima,Valle), Ecuador
(Azuay, Carchi, Chimborazo, Cuenca, Imbabura, Loja, Morona-Santiago, Napo, Specimens
examined: COLOMBIA. Cauca: Carretera a La Plata, de Puracé al Alto de San Rafael, GarciaBarriga & Hawkes 12833 (LIL, US); Cordillera Central, Puracé, Sneidern 1781 (F, GH, NY),
Sneidern 1782 (K, S), Barclay & Schultes 155 (GH); Páramo de Moras, between Mozoco and
Pitayo, Pittier 1513 (NY). Mariquita: Tolima, pres du Boqueron, Linden 924 (G, P). Nariño:
Zona fría, norte de Yacuanquer, Garganta 465 (F). Putumayo: río Putumayo, filo de la Cordillera
enter El Encano y Sibundoy, páramo de San Antonio del Bodoncillo, Cuatrecasas 11751 (F).
Quindio: Mpio Pijao, Páramo de Chili, above Finca Maizopolis, 4°15-20′N, 75°35′W, Juteyn et
al. 12951 (GH, NY); Mpio Salento, Verde Cocóra, 4°40-45′N, 75°20-25′W, Luteyn et al. 13025
(GH, NY). Tolima: Rosalito, near Páramo de Ruiz, Pennell 2960 (GH, NY). Valle: Cordiller
Central, cabeceras de los ríos Tuluá y Bugalagrande, Páramo de Las Vegas, Cuatrecasas 20309
(F). ECUADOR. Laguna Cocha, André 3020 (GH, K, NY). Azuay: Gualaceo-Limón rd, km
25.2, 78°39-40′W 03°00′S, Jørgensen et al. 1853 (MO); Páramo del Castillo, Camp E-5125 (GH,
NY); Páramo de Tinajillas, 30-50 km S Cuenca, Camp E-2267 (GH, NY). Carchi: Páramo de El
Angel. Laguna Sur de El Voladero, 78°26′W 00°37′S, León 1083 (AAU); road from El Playon
de San Francisco to Cerro Mirador, 77°39′W, 00°36′S, Øllgaard 98159 (AAU); TulcánMaldonado hwy, km 26 Cueva de Botas, Boeke 797 (SI); Páramos de Angel, Barclay & Juajibioy
9405 (GH, MO), Benoist 4626 (P). Chimborazo: Camino de Pusucucho al Placer, Acosta Solís
7272 (F), Acosta Solís 7291 (F); Páramo de Carnicería, faldas volcán Sangay, Acosta Solís 7677
(F). Cotopaxi-Tungurahua: Llanganates, Aucacocha, laguna Aucacocha, Jaramillo 6021 (MO).
Cuenca: Cerro Yanghuang, near Pindilic, Lehmann 5604 (F, GH, K). Imbabura: Lago San
Marcos, Cayambe, Cazalet & Pennington 5313 (B, K, NY, UC, US); near Laguna Yanuyacu,
Wiggins 10431 (DS); E slopes of Cayambe Peak, Wiggins 10409 (DS, GH); N slopes of
Cayambe on headwaters of E fork of Río Desaguadero, Wiggins 10385 (DS). Loja: Cordillera de
Las Lagunitas, Amaluza-Jimbura-Zumba, Km 39.2 at pass, 79°25′22″W, 4°44′38″S, Jørgensen
et al. 938 (LOJA, MO, QCA, QCNE); between Tambo Cachiyacu, La Entrada and Nudo de
Sabanillas, Steyermark 54459 (F, GH, NY); Parque Nacional Rodocarpus, road from Yangana to
61
radio towers on Cerro Toledo, van der Werff & Palacios 9182 (MO); Yangana-Cerro Toledo
road, 70°6′W, 4°23′S, Øllgaard et al. 58238 (AAU); E of Nudo de Cajanuma, 79°10′W, 4°05′S,
Øllgaard 74675 (AAU); 14 km E Loja, 4°0′S, 79°1′W, Knight 910 (GH). Morona-Santiago:
Parque Nacional Sangay, between Plazapamba and Yanayacu, 78°25′W, 02°02′S, Clark et al.
1828 (MO, QCNE); between Culebrillas and Alao, 78°30′W, 02°00′S, Clark et al. 1876 (MO,
QCNE); Cuenca_General Plaza (Limón), km 29, Sparre 18786 (S). Napo: Volcán Cayambe,
77°59′W, 00°3′S, Øllgaard et al. 34271 (MO); Quijos Cantón, Reserva Ecológica Antisana,
Carretere Pifo-Baeza, Páramo de la Virgen, 78°12′W, 00°23′S, Freire-Fierro, Vargas & Narváez
2874 (MO). Pichincha: rd from Olmedo to Laguna San Marcos, N slope of Volcán Cayambe,
0°7′N, 78°0′W, Balslev & Steere 4474 (NY). Tungurahua: 2.5 km W of Cerro Hermoso,
1°13′S, 78°18′W, Holm-Nielsen & Jaramillo 28656 (AAU, NY); upper slopes of Volcano
Tungurahua, Popenoe 1286 (NA); Páramo Minza, Penland & Summers 327 (DS, F, GH);
Santiago de Pillaro, Parque nacional Llanganates, Camino desde el Valle de los Frailejones hasta
el pamo de Soguillas, 78°16′W, 01°08′S, Narváez et al. 523 (MO); Laguna Soquillas, Clark &
Fair 3508 (MO, QCNE). VENEZUELA. Merida: Laguna Mucubaji, above Los Aparaderos,
Steyermark 57507 (F, GH); Sierra de Santo Dominigo, Laguna Negra, Dennis 2147 (K).
Trujillo-Merida: Andes, Jameson 179 (G).
Schulz (1903) treated some Venezuelan and Colombian plants of Cardamine jamesonii as members
of the African C. johnstonii (as pro. punicea and var. superba, respectively). He also treated the latter
species as distinct from the African C. oblique A. Richard, which he treated as occurring both in Africa
and Mexico. Baehni (1937) disagreed with such a discrepancy and recognized C. punicea as a species
distinct form the African C. johnstonii. However, he overlooked the fact that C. punicea is
indistinguishable from C. jamesonii. In my opinion, the alleged trans-Atlantic distributions by Schulz
(1903) represent misidentifications of native New World taxa superficially resembling some of the
African species.
The variability in leaflets morphology and flower color has lead to the recognition of several taxa
within Cardamine jamesonii. However, the variation is continuous in every aspect of the plant, and it is
impractical to divide the species into infraspecific taxa. The only exception is the type collection of C.
tolimensis, which differs from the other collections of C. jamesonii by being densely pubescent
throughout and by having six pairs of leaflets. It might represent an inDep. endent taxon, but for now it
is tentatively placed in C. jamesonii.
Cardamine jamesonii is readily distinguished from C. ovata by having purple, violet, red, or lavender
(vs. white) flowers, larger petals ((8−)10−14 × 5−8 mm (vs. (−)− × − mm) and middle stem leaves with
3−5(−6) pairs (vs. (1−)2(−4) pairs) of lateral leaflets. From C. lojanensis, C. jamesonii is readily
distinguished by having purple, violet, red, or lavender (vs. white) flowers, non-tuberous and non-scaly
(vs. tuberous and scaly) rhizomes, and cauline leaves with 3−5(−6) pairs (vs. 6−12 pairs) of lateral
leaflets.
Pichincha,Tungurahua), Vezuela (Merida, Trujillo).
Cardamine jejuna Standley & Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23: 54. 1944.
Holotype: Guatemala, J. Steyermark 50569 (holotype, F!).
Herbs, annual, sometimes scapose. Stems 5–10 cm, erect, simple, leafless or 1-leaved,
glabrous throughout. Basal leaves 1.5–3 cm, rosulate, pinnately compound, petiolate; terminal
lobe 3–6 × 3–6 mm, suborbicular to broadly ovate, obscurely 3-lobed and the lobes mucronate,
glabrous or sparsely hirsute adaxially with whitish flattened trichomes, base truncate to rounded,
62
margin entire, apex obtuse to rounded, long petiolulate; lateral lobes 1 or 2 each side of midvein,
short petiolulate, similar to terminal one. Raceme ebracteate, 3–5-flowered, lax; fruiting pedicels
1–1.5 cm, erect, slender. Sepals 1.5–2 × ca. 0.7 mm, oblong; petals 3–4 × 1–2 mm, white,
obovate; filaments 2–2.5 mm; anthers ca. 0.5 mm; ovules ca. 14 per ovary. Fruits 2–2.5 cm ×
0.7–1 mm; style ca. 1.5 mm. Seeds ca. 1.5 × 0.6 mm, oblong.
Flowering: Aug.
Habitat: forests.
Elevation: ca. 2800 m.
Distribution: Mexico (Chiapas), Costa Rica.
Cardamine jonselliana Al-Shehbaz, Harvard Pap. Bot. 9: 5. 2004. TYPE: Uganda, Ruwenzori,
Stuhlmann Pass, on moist open ground amongst grasses, 13,700 ft, between 18 Jun and 5 Jul
1968, A. C. Hamilton 742 (holotype, K!).
Herbs, perennial, glbrous throughout except for leaf margin. Rhizomes branched, not scaly,
3–5 mm wide. Stems 10–25 cm, erect, simple, usually rooting from lower nodes. Rhizomal
leaves and basal absent; cauline leaves pinnately compound, 5–9-foliolate; petiole 2–4 cm, not
auriculate at base; terminal leaflet ovate to orbicular, 4–10 mm, obscurely 3–5-lobed, repand,
ciliate, on a petiolule 1–2 mm; lateral leaflets orbicular to broadly ovate, 2–6 mm, sessile, repand
to entire. Raceme bracteate throughout, 12–25-flowered; lowermost bracts 5-foliolate, gradually
reduced in size and number of leaflets upward, uppermost repand or obscurely lobed; fruiting
pedicels ascending to suberect, straight, stout, 5–10 mm. Sepals reddish purple, oblong, 3–3.5
mm, glabrous; petals purple, spatulate, 7–9 × 3–3.5 mm, obtuse at apex, not differentiated into
blade and claw; stamens 6, filaments 2.5–3.5 mm, anthers oblong, 0.7–1 mm. Fruits (young) 1–
1.5 cm. Seeds not seen.
Flowering Jun–Jul.
Habitat: moist and open grounds among grasses.
Elevation: ca. 4170 m.
Distribution: Uganda.
Cardamine keysseri O. E. Schulz, Bot. Jahrb. Syst. 62: 480. 1929. TYPE:
Herbs annual or short-lived perennial. Rhizome slender, prostrate. Stems 1–4 dm, erect,
rooting from lower nodes, hispidulous basally. Leaves pinnately compound, 5–9-foliolate,
glabrous or sparsely pubescent; basal leaves 4–8 cm, petiole 1–4 cm, soon withered; cauline
leaves to 9 cm, with non-auriculate petioles; leaflets trifid, 0.3–2.5 × 0.2–1.4 cm, base attenualte
to cuneate or rarely truncate, apex acute; terminal leaflet larger, sessile or with petiolule to 1.5
cm. Raceme ebracteate, 5–10-flowered; fruiting pedicels 4–10 mm, divaricate. Sepals oblong,
2.5–3.5 mm; petals white, spatulate, 7–8 × 2.5–3.5 mm, obtuse or emarginate; stamens 6,
subequal, filaments subulate; anthers ca. 0.8 mm; ovules – per ovary. Fruits linear, 2–3.8 × 1–2
mm; style 0.5–1.4 mm. Seeds elliptic, 1.4–1.8 × 1.2–1.4 mm, smooth.
Habitat: Sandy or gravelly streamsides, forest clearings.
Elevation: 900–4000 m.
Distribution: New Guinea.
Cardamine kitaibellii Becherer, Ber. Schweiz. Bot. Ges. 43: 57. 1934; Dentaria polyphylla
Waldstein & Kitaibel, Descr. Icon. Pl. Hung. 2: 174, t. 160. 1805, not C. polyphylla D. Don, nor
O. E. Schulz. TYPE: E Croatiae montibus ad Koreniczam et Priboy, Herb. Kitaibel, no. LI/111
(lectotype designated by Kováts (Ann. Hist.-Nat. Mus. Natl. Hung. 84: 41. 1992).
63
Cardamine komarovii Nakai, Repert. Spec. Nov. Regni Veg. 13: 271. 1914; Alliaria auriculata
Komarov, Trudy Imp. S.-Peterburgsk. Bot. Sada 18: 437. 1901, not Cardamine auriculata S.
Watson (1882).
Arabis cebennensis de Candolle var. coreana H. Léveillé, Bull. Acad. Géog. Bot. 19: 260. 1909.
TYPE: Korea. “In petrosis rivorum montis des diamants,” 24 June 1906, Faurie 570
(holotype, E!).
Herbs, perennial, 12–75 cm. Rhizomes short, stout. Stems erect, simple at base, branched
above, finely striate, pubescent or glabrescent. Basal leaves rosulate; petiole 2–5 cm, winged at
base, glabrous or pilose; leaf blade broadly cordate or suborbicular, 1.5–4 × 1.5–3.5 cm, margin
coarsely dentate, the teeth submucronate; cauline leaves 3–8, simple; petiole winged, (0.5–)1–4
cm, auriculate-amplexicaul at base; auricles oblong or lanceolate, 2–10 × 0.5–4 mm, entire; leaf
blade cordate or broadly ovate, (2–)3–7.5(–9) × (1.2–)2.5–6 cm, pilose at least along veins and
margins, rarely glabrous, base cordate, margin coarsely and irregularly toothed, sometimes
incised on distal leaves, apex acuminate or obtuse. Racemes ebracteate; fruiting pedicels
divaricate or ascending, 1–2.5 cm, slender, pilose or glabrous. Sepals oblong, 2.5–3 × 1.2–1.6
mm, glabrous or pilose, base not saccate; petals white, broadly obovate, 3–4 × ca. 1.5 mm, apex
rounded; filaments 2–3 mm; anthers oblong, ca. 1.5 mm; ovules 6–12 per ovary. Fruit linear, 2–
4 cm × ca. 1 mm; gynophore stout, 0.2–0.7 mm; valves glabrous or sparsely pilose, not torulose,
acuminate; style 2–3 mm, slender. Seeds brown, oblong, 1.5–2 × 1–2 mm.
Flowering: May–Jun.
Habitat: streamsides, moist areas.
Elevation: 700–1000 m.
Distribution: China (Heilongjiang, Jilin, Liaoning), Korea.
Cardamine kruesselii Johow, Stud. Fl. Isals Juan Fernandez 112. 1896. TYPE: Chile. Juan
Fernandez Islands: Masafuera, Johow s.n. (holotype,?)
Herbs, annual? perennial?, densely hirsute throughout. Rhizome tuberous?. Stems 5–15 cm,
densely hirsute, subangular, branched near base and above, subdecumbent or ascending, flexous
above. Basal leaves not seen; cauline leaves 3–5-foliolate, 2–5 cm; petiole hirsute, 1–2 cm;
terminal leaflet 8–15 × 1–6 mm, entire or subapically toothed, ovate or obovate to narrowly
lanceolate, subsessile or on a petiolule to 3 mm; lateral leaflets oblong to ovate or narrowly
lanceolate, 4–10 × 1–7 mm, entire, cuneate or oblique at base. Raceme ebracteate or lowermost 1
or two flowers bracteate; rachis flexuous; fruiting pedicels slender, divaricate, 3–6.5 mm. Sepals
oblong, glabrous, 1–1.5 × 0.3–0.7 mm; petals white, spatulate, 2.5–3 × 0.3–0.8.1 mm; stamens #
?; filaments 1–1.5 mm; anthers ovate, 0.1–0.2 mm; ovules 18–22 per ovary. Fruit linear, 1–2.5
cm × 0.8–1 mm, replum sparsely pubescent; style 1–1.5 mm. Seeds light brown, ovate, 0.9–1.2 ×
0.6–0.7 mm, wingless.
Flowering: Dec.
Distribution: Chile (Juan Fernandez Islands/Mas Afuera).
Specimens examined: CHILE. Juan Fernandez Islands: Mas Afuera, [500 m], [33°45′S,
80°46′W], 28 Dec. 1891, F. Johow s.n. (CONC).
An obscurely known species for which I have seen only the specimen above. It is said to be a
perennial, but the material examined is fragmentary. The presence of racemes bracteate basally
and the rhizome suggest affinity to Cardamine bonariensis, if indeed different. However, the
smaller size of flowers and the pubescent replum argue against that. Schulz (1903) indicated
affinity to C. hispidula, but I have not seen material of the latter.
64
Cardamine lacustris (Garnock-Jones & P. N. Johnson) Heenan, New Zealand J. Bot. 40: 568.
2002; Iti lacustris Garnock-Jones & P. N. Johnson, New Zealnd J. Bot. 25: 603. 1987. TYPE:
New Zealand, Henry Creek, E side of Lake Te Anau, 21 May 1971, P. N. Johnson s.n.
(holotype CHR 286399).
Herbs, annual or ephemeral, glabrous or sparsely pubescent. Stems to 5 cm, prostrate,
geniculate. Basal leaves petiolate, rosulate, early ones entire, linear-spatulate; cauline leaves and
later basal ones pinnatifid, 2–3(–5) cm; petiole 2–12 mm, not auriculate at base; terminal lobe
narrowly spatulate, 2–3(–5) mm wide; lateral lobes 1 or 1(–3) on each side, narrowly oblongspatulate, 4–10 mm. Inflorescence solitary flowers or 2(or 3) from leaf axils; fruiting pedicels 2–
5 mm. Sepals oblong, suberect, glabrous, 1.5–2 mm; petals white, suberect, oblong- to obovatespatulate, 1.8–2.5 × 0.8–1.2 mm; stamens 4(or 6), median pair(s) 1.6–2 mm, lateral 1–1.5 mm;
ovules 10–16 per ovary. Fruits ellipsoid to oblong, (1–)2–3(–3.5) × 1–1.7 mm, convex, glabrous.
Seeds oblong, ca. 1 mm. 2n = 48.
Flowering: Mar–May.
Habitat: lake shores.
Distribution: New Zealand (South Island).
Cardamine latior Heenan, New Zealand J. Bot. 46: 562. 2008. TYPE: New Zealand, Auckland
Islands, Bivouac Saddle, fellfied, 1600′, Jan 1963, E. J. Godley s.n. (holotype, CHR 134244A;
isotype, CHR).
Herbs, perennial, cespitose. Basal leaves compactly rosulate, pinnately compound, 11–15foliolate, to 4(–5) cm; petioles 8–12 mm; leaflets often overlapping towards leaf apex, reniform
to orbicular or broadly elliptic, entire; terminal leaflet 3.5–4.7 × 3.5–4.1 mm, with 2 lateral lobes;
lateral leaflets 1.5–2.5 × 0.7–2.2 mm; cauline leaves sometimes present, not auriculate at base,
similar to basal leaves but smaller and with fewer leaflets. Racemes basally bracteate, not
elongated in fruit; fruiting pedicels 5–10(–17) mm, erect. Sepals 2–2.4 mm, oblong to elliptic,
glabrous; petals white to pale pink or purple, 3.2–4 × 1.2–1.5 mm, obovate, apex obutse to
rounded; stamens 6, filaments 1.6–2.4 mm, anthers 0.4–0.5 mm. Fruits 1.2–2.3 cm × 1.7–2.2
mm, glabrous; style 0.7–1.4 mm. Seeds 1.4–2.1 × 1.1–1.5 mm, rounded to oblong.
Flowering: Oct–Dec.
Habitat: fellfields, among boulders, scree.
Distribution: New Zealand (Auckland and Adams islands).
Cardamine lazica Boissier & Balansa in Boissier, Fl. Or. Suppl. 31. 1888; Cardamine amara
Linnaeus prol. Lazica (Boissier & Blansa) O. E. Schulz, Bot. Jahrb. Syst. 32: 500. 1903.
TYPE: Turkey, in Ponto litorali ad rivulos prope Rize, 100m, et in regione subalpina inter
Andon et Djimil, 1850, 1866, Balansa s.n. (lectotype, needs designation)
Cardamine leucantha (Tausch) O. E. Schulz, Bot. Jahrb. Syst. 32: 403. 1903; Dentaria
leucantha Tausch, Flora 19: 404. 1836. TYPE:
Cardamine cathayensis Migo, J. Shanghai Sci. Inst. Sect. III. 3: 223. 1937.
Cardamine koreana (Nakai) Nakai, Bull. Nat. Sci. Mus. Tokyo 31: 49. 1952. List bsionym.
Cardamine leucantha (Tausch) O. E. Schulz var. crenata D. C. Zhang, Bull. Bot. Res., Harbin
11: 42. 1991. TYPE: China, Anhui, Jingdexian, Yucun Wolonggou, 200 m, 7 May 1985,
Shao Jianzhang 82143 (holotype, ANU).
Cardamine leucantha (Tausch) O. E. Schulz var. glaberrima F. Maekawa ex H. Hara, J. Jap.
Bot. 51: 192. 1976.
65
Cardamine macrophylla Willdenow var. albiflora Makino, Bot. Mag. (Tokyo) 8: 302. 1894.
Cardamine macrophylla Willdenow var. parviflora Trautvetter & C. A. Meyer, Fl. Ochot. 15
1856.
Dentaria dasyloba Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 27: 296. 1854; Cardamine
dasyloba (Turczaninow) Miquel, Ann. Mus. Bot. Lugduno-Batavi 2: 73. 1866; D.
macrophylla (Willdenow) Bunge ex Maximowicz var. dasyloba (Turczaninow) Makino, Bot.
Mag. (Tokyo) 11: 157. 1897. TYPE:
Herbs, perennial, sparsely to densely villous. Rhizomes creeping, slender, not scaly or
stoloniferous. Stems 2.5–7.5 dm, simple, flexuous. Cauline leaves 4–7, including petiole
(8–)10–20(–25) cm; petiole (1–)2–8(–10) cm, not auriculate at base; terminal leaflet lanceolate,
elliptic, to ovate-elliptic, (3–)4–9(–13) × (0.6–)1–3.5(–4) cm, adaxially shortly strigose or
puberulent, abaxially pilose or long strigose, with a petiolule 5–13(–20) mm, base cuneate,
margin antrorsely ciliate and irregularly serrate, double serrate, or subdentate, apex acuminate or
rarely acute; lateral leaflets 2 or 3 pairs, rarely uppermost leaf trifoliolate, similar to terminal one,
smaller, sessile, base cuneate or sometimes oblique. Racemes ebracteate, 12–24-flowered;
fruiting pedicels divaricate or ascending, (0.5–)1–1.8(–2.3) cm, pilose or puberulent, slender.
Sepals oblong, (2–)2.5–3.5 × 0.9–1.5 mm, margin membranous, pilose abaxially; petals white,
spatulate to oblong-oblanceolate, 6–8 × 2–3.5(–4) mm, cuneate and not clawed at base, apex
rounded; median filament pairs 5–6 mm, lateral pair 4–5 mm; anther oblong, 0.8–1.3 mm; ovules
6–12 per ovary. Fruit linear, (1–)1.5–3 cm × 1–1.5 mm; gynophore 0.3–0.8(–1) mm; valves
smooth, sparsely hairy or glabrous; style slender, (2–)3–5 mm. Seeds brown, oblong, 1.5–2.2 ×
0.8–1.2 mm, narrowly winged or wingless. 2n = 16.
Flowering: Apr–Jul.
Habitat: roadsides, shady areas, forests, wet places along streams, roadsides.
Elevation: 100–2000 m.
Distribution: China (Anhui, Gansu, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Jiangxi, Jilin,
Jiangsu, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shanxi, Sichuan), Japan (Honshu, Hokkaido),
Korea, Mongolia, Russia (Far East).
Cardamine lihengiana Al-Shehbaz, Novon 10: 326. 2000. TYPE: China. Yunnan. Yiliang, Xia
Cao Ba, 1900 m, 16 Jun 1982, Li Heng, Chen Yü & Yü Hongyuan 1267 (holotype, KUN!;
isotype, KUN!).
Herbs, perennial, glabrous throughout. Rhizomes slender, without stolons. Stems 3–5 dm,
erect, angled, with internodes usually more than 2 cm. Rhizomal leaves and lower cauline
simple; petiole 1–4 cm, not auriculate at base; leaf blade reniform or suborbicular, 0.7–2.5 × 1–3
cm, palmately veined, base cordate, margin repand-crenate, obscurely to distinctly 5- or 7-angled,
apex obtuse, obscurely mucronulate; middle and upper cauline leaves with axillary flowers,
simple or rarely 1 or 2 with a lateral, leafletlike lobe, suborbicular or ovate, angled, not crenate,
progressively smaller upward. Racemes bracteate throughout, very lax; fruiting pedicels
ascending, 1.5–3 cm, slender, straight. Sepals ovate, 1.5–1.8 × 0.9–1 mm, apex membranous,
base not saccate; petals white, obovate, 3.5–4.5 × 1.5–2 mm, not clawed, apex rounded; median
filament pairs ca. 2.5 mm, lateral pair ca. 2 mm; anthers oblong, ca. 0.6 mm; ovules 20–24 per
ovary. Fruit linear, 2–3.6 cm × ca. 1 mm, sessile; valves glabrous, smooth; style 1–2 mm. Seeds
brown, oblong, 1.2–1.4 × 0.7–0.9 mm, wingless.
Flowering May–Jun.
Elevation: ca. 1900 m.
66
Distribution: China (Yunnan).
Cardamine lilacina W. J. Hooker, Comp. Bot. Mag. 1: 273. 1835; C. pratensis var. lilacina (W.
J. Hooker) J. D. Hooker, Fl. Tasmaniae 1: 19. 1855. TYPE: Australia, banks of the Macquarie
River, C. Fraser s.n. (lectotype here designated, K).
Herbs, annual or perennial, sometimes forming cushions, often glabrous. Rhizomes present.
Stems to 5 dm, erect to decumbent. Basal leaves pinnately compound, with 1–3 pairs of lateral
leaflets; leaflets often ovate, to lanceolate, undivided, entire; cauline leaves petiolate, not
auriculate at base, less divided, reduced in size upwards; uppermost lanceolate, entire. Raceme
ebracteate, corymbose, elongated in fruit; fruiting pedicels erect to spreading, 5–20 mm, slender.
Sepals 2–3 mm, purple, obling; petals 7–11 mm, pink to purple; stamens 6; ovules –. Fruit
linear, 1–4.5 cm × 1–2 mm; style 1–3 mm. Seeds oblong to ovoid, 1.5–2.5 mm, flattened,
narrowly winged.
Habitat: damp forests, wet grounds.
Distribution: Australia (New South Wales, Tasmania, Victoria).
Cardamine lineariloba I. Thompson, Muelleria 9: 163. 1996. TYPE: Australia, Victoria, Mt.
Arapiles, SE slope along watercourse, ca. 1.6 km down from top, 22 Sep 1968, A. C.
Beauglehole 28699 (holotype, MEL).
Herbs, annual, glabrous throughout. Stems to 25 cm, erect to ascending, usually much
branched from base and above. Basal leaves not fleshy, not rosulate, petiolate, to 6 cm, simple,
narrowly oblanceolate, entire, base attenuate, sometimes pinnatisect and with 1−several, linear
lateral lobes much smaller than the sublinear terminal lobe to 2 cm; cauline leaves several,
similar to basal, petiolate, not auriculate at base, gradually reduced in size upwards. Racemes
ebracteate, often few flowered; fruiting pedicels divaricate-ascending, 5−10 mm. Sepals ovate,
1.5−2 mm; petals white, cuneate, 3−5 mm; stamens 6. Fruits linear, erect, 0.9−2 cm × ca. 1 mm;
style to 1 mm. Seeds elliptic, to 1.2 mm.
Flowering: Aug−Sep.
Habitat: lowland areas along stream banks, seasonally wet lowlands, swamps.
Elevation:
Distribution: Australia (South Australia, Victoria).
Cardamine lojanensis Al-Shehbaz, Novon 7: 6. 1997. TYPE: Ecuador, Loja, Parque nacional
Podocarpus, Cerro Toledo E of Yangana, wet paramo around radio station, 3400−3450 m,
4°24′S, 76°6′W, 26 Feb 1985, B. Øllgaard, S. Laegaard, K. Thomsen, K. Korning & T. Illum
58147 (holotype, AAU!).
Herbs, perennial. Rhizomes tuberous, scaly, with persistent expanded petiolar bases of
previous years. Stems 1 or 2, 15−40 cm, erect, somewhat striate, glabrous or sparsely puberulent.
Basal leaves compound, with 13−23 leaflets; petiole 1.5−9 cm, 1−3× as long as leaf rachis,
distinctly expanded into a triangular, winged, persistent base becoming scalelike in subsequent
seasons; rachis grooved, 0.7−4 cm; lateral leaflets petiolulate, ovate or ovate-oblong to lanceolate
in outline, 2−9 × 1−7 mm, pinnatifid, laciniate, to incised, the segments incised to dentate,
margin sparsely ciliate with minute trichomes 0.06−0.12 mm, apex acute; petiolule grooved,
(0.5−)1−3 mm; cauline leaves 2−6, with 13−19 leaflets, similar in morphology to basal leaves,
reduced in size and division of leaflets upward;p petiole 0.7−1.2 cm; leaflets short petiolulate to
subsessile in uppermost leaves. Racemes basally bracteates, corymbose, elongated considerably
67
in fruit; bracts adnate to pedicel; fruiting pedicels ascending, straight, glabrous, striate,
lowermost 2−2.5 cm. Sepals oblong, 3−4 × 1.2−1.5 mm, slightly saccate, erect or suberect,
caducous, glabrous; petals white, broadly obovate to suborbicular, 8−11 × 6−7 mm, abrubtly
narrowed into a clawlike base to 1 mm, apex shallowly emarginate to subtruncate; filaments
white, 3−5 mm; anthers oblong, ca. 1 mm. Fruits linear, 2.5−5 cm × ca. 2 mm; gynophore ca. 1
mm; style 3−4 mm. Seeds brown, oblong, ca. 2.4 × 1.2 mm.
Flowering: Nov−Feb.
Habitat: sandy rocky soil, wet paramo.
Elevation: 3150−3500 m
Distribution: Ecuador (Loja).
Specimens examined: ECUADOR. Loja: road from Yangana to Cerro Toledo, km 18−22 to the
antennas, 79°6′40−42″W, 4°22′28″−4°24′13″S, Jørgensen, Ulloa & Caranqui 2179 (LOJA, MO,
QCA, QCNE).
Cardamine longii Fernald, Rhodora 19: 91. 1917. TYPE:
Herbs, perennial, glabrous throughout. Rhizomes slender, cylindrical, 0.8–1.5 mm in diam.;
stolons absent. Stems 0.5–4(–6) dm, prostrate to decumbent or erect, simple or branched,
glabrous. Rhizomal leaves absent; cauline leaves 3–10, simple or rarely 3–5-foliolate, somewhat
fleshy; petiole 0.4–2.5 cm, base not auriculate; leaf blade or terminal leaflet of compound leaves
orbicular to reniform or ovate to oblong, 0.4–3 × 0.3–2 cm, base subcordate to rounded or
subtruncat, margin entire or repand or rarely undulate, apex rounded; lateral leaflets (when
present) similar to terminal one but considerably smaller, sessile or with a petiolule to 2 mm.
Racemes ebracteate; fruiting pedicels divaricate-ascending to spreading, 0.5–2(–4) mm. Sepals
ovate, 0.7–1.2 × 0.5–0.7 mm, erect, base of lateral pair not saccate; petals absent or rudimenatary
and to 0.7 mm; filaments 0.5–0.8 mm; anthers ovate, 0.1–0.2 mm. Fruit narrowly oblong to
linear, (3–)5–10(–15) × 0.8–1.2 mm; valves glabrous; style 0.2–0.5(–1) mm; ovules and seeds 6–
22 per fruit. Seeds light or yellowish brown, oblong or ovate, 1–1.4 × 0.7–1 mm.
Flowering: Jun–Sep.
Habitat: tidal marshes, mud flats, tidal shores of rivers, shallow water, swampy areas, shady
rocky crevices covered at high tide.
Elevation: sea level.
Distribution: United States (Florida, Maine, Maryland, Massachusetts, New Jersey, New York,
North Carolina, South Carolina, Virgina).
Cardamine longipedicellata Rollins, J. Arnold Arbor. 21: 395. 1940. TYPE: Mexico, Nuevo
Leon, San Francisco Canyon, about 15 miles SW of Pueblo Galeana, 11 May 1934, C.H. &
M.T. Mueller 298 (holotype, GH!).
Cardamine ramosa Rollins, Cruciferae Continental N. Amer. 295. 1993. TYPE: Mexico, San
Luis Potosí, 3 miles E of Xilitla, 3000 ft, 18 Jul 1963, R. L. McGregor, L. J. Harms, A. J.
Robinson, R. del Rosario & R. Segal 903 (holotype, GH!).
Herbs, annual or rhizomatous perennial, glabrous or sparsely pubescent. Stems 10–30 cm,
erect or decumbent, simple or branched above. Cauline leaves petiolate, not auriculate, pinnately
compound; terminal leaflet 5–20 × 3–8 mm, suborbicular, broadly ovate, or obovate, base
cuneate to subtruncate, margin entire or dentate, apex acute, petiolule to 2 cm; lateral leaflets 1–
3(or 4) on each side of rachis, similar to terminal leaflet but smaller and often with shorter
petiolules; uppermost leaves smaller. Racemes ebracteate, lax, few flowered, somewhat secund
in fruit; fruiting pedicels (4–)9–20 mm, divaricate to ascending, slender, straight, glabrous or
68
pubescent. Sepals 2–2.5 × 0.8–1 mm, oblong, caducous; petals 4–6 × 1–2 mm, white, spatulate,
apex obtuse; filaments 2–3 mm; anthers 0.6–0.7 mm, oblong; ovules 12–16(–18) per ovary.
Fruits 2.2–3 cm × 1–1.3 mm; style 0.5–2 mm. Seeds 1.2–1.5 × 0.8–1 mm, light brown, oblong.
Flowering: Jul–Nov.
Habitat: slopes, roadsides.
Elevation: 1000–3200 m.
Distribution: Mexico (Chiapas, Hidalgo, Distito Federal, Nuevo León, Puebla, San Luis Potosí,
Veracruz), Guatemala.
Notes: the only difference in the original descriptions of Cardamine longipedicellata and C.
ramosa is in the length of the fruiting pedicels, which was said to be 10-20 mm long in the
former and 4-7(-10) mm in the latter. However, several paratypes cited by Rollins (1993)
under C. ramosa, including Matuda 4694, clearly have fruiting pedicels to 17 mm.
Cardamine loxostemonoides O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 9: 1069. 1926;
Loxostemon loxostemonoides (O. E. Schulz) Y. C. Lan & T. Y. Cheo, Bull. Bot. Res., Harbin
1(3): 54. 1981. TYPE: NW India, Nipchang valley, 4300–4600 m, 31 Aug 1884, Duthie 2724
(lectotype, designated by Rashid & Ohba (1993), ?B; isolectotype, K, DD).
Loxostemon incanus R. C. Fang ex T. Y. Cheo & Y. C. Lan, Bull. Bot. Res., Harbin 1(3): 54.
1981. TYPE: China, Yunnan, Chungtien Plateau, 7 Jul 1930, K. M. Feng 1559 (holotype,
KUN).
Cardamine tibetana Rashid & H. Ohba, J. Jap. Bot. 68: 206. 1993. TYPE: China, SE Tibet, Tse
La, Langong, 94°00′n, 28°45′E, 14–15000 ft, open bare scree, 21 Jun 1938, F. Ludlow & G.
Sherriff 5618 (holotype, TI!; isotypes, BM!, E!).
Herbs, perennial, glabrous or sparsely to densely pilose. Rhizomes slender, with several
bulbils and stolons; bulbils with fleshy, white, scaly leaves apically with rudimentary
appendages. Stems (5−)12−30(−35) cm, somewhat decumbent and slender below, simple.
Rhizomal leaves glabrous or pilose, compound; petiole (0.7−)2−15(−20) cm; terminal leaflet
with a petiolule 3−10 mm, blade undivided and suborbicular, oblanceolate, or linear, or trifid and
suborbicular to broadly obovate in outline and with obovate or oblong lobes the basal pair of
which sometimes with a tiny lobule, terminal lobe 0.6−2(−3) cm × 2−5 mm; lateral leaflets 2−5
pairs, sessile or petiolulate, similar in shape and division to terminal lobe but smaller. Cauline
leaves 1−4; petiole (0.3−)1−3(−4) cm, not auriculate at base; terminal and lateral leaflets similar
in shape, size, and number to those of basal leaves. Racemes ebracteate, 2−14-flowered; fruiting
pedicels (5−)1−2.5 cm, ascending to suberect, glabrous. Sepals broadly ovate, 2.5−5 × 1.5−2.5
mm, glabrous, broadly membranous at margin and apex, base of lateral pair subsaccate; petals
purple with darker veins, broadly obovate, 0.8−1.2(−1.4) cm × 5−8(−8.5) mm, not clawed, apex
rounded; median filament pairs (3.5−)4.5−6 mm, lateral pair 2.5−4 mm; anthers narrowly oblong,
1.4−2 mm; ovules 14−20 per ovary. Fruits linear, 2.5−3.5 cm × 1.2−1.5 mm; valves glabrous;
style 1−3 mm. Seeds brown, ovate, ca. 1.5 × 1 mm, wingless.
Flowering: Jun−Jul.
Habitat: mountains slopes, along ditches, damp grounds by streams, open grass and gravel, scree.
Elevation: 2900–5500 m.
Distribution: Bhutan, China (Xizang), India (Sikkim), Kashmir, Nepal.
Cardamine luxurians (O. E. Schulz) Rashid & H. Ohba, J. Jap. Bot. 68: 205. 1993; C.
loxostemonoides O. E. Schulz var. luxurians O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 9:
69
1070. 1927. TYPE: NW India, Tihri-Garwal, Nila Valley, 16 Aug 1883, Duthie 912 (holotype,
B!; isotype, K!).
Herbs, perennial. Rhizomes branched, stoloniferous, nontuberous, with membranous scales.
Stems 11−25 cm; erect to procumbent, glabours. Basal leaves and cauline leaves similar, with
5−11 leaflets; petiole 2−9 cm; lateral leaflets on petiolules 2−10 mm, broadly ovate to
suborbicular, 13−18 mm, 6−13 mm wide, base cuneate to oblique, margin subentire or 3-toothed
or lobed, apex suacute and minutely mucronate; terminal leaflet slightly larger. Racemes
ebracteate, 6−20-flowered; fruiting pedicels 1−2 cm, straight, ascending. Sepals 3−3.7 mm,
1.5−2.2 mm wide, oblong; petals purplish to pink with darker veins, obovate, 10−14 mm, 4−6
mm wide; median filaments 6−7.5 mm, lateral filaments 3.5−4 mm; anthers oblong, 0.9−1.2 mm;
ovules 16−20. Fruits linear, 2.5−4.5 cm, 1−1.5 mm wide; valves glabrous; style ca. 1 mm. Seeds
not seen. 2n = 32.
Habitat: shade of boulders.
Elevation: ca. 3350 m.
Distibution: Kashmir.
Cardamine lyrata Bunge, Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2: 29. 1833.
Cardamine argyi H. Léveillé, Mem. Real Acad. Cienc. Art., Barcelona 12: 547. 1916. TYPE.
China, Jiangsu, without locality and collection date, d’Argy s.n. (holotype, E).
Herbs, perennial, glabrous throughout. Rhizomes short, thick, not scaly; stolons from
rhizome or lower to middle stem nodes to 80 cm. Stems (2–)3–7(–8) dm tall erect, simple, striate
angled, 8–15-leaved. Leaves on stolons simple or rarely with a basal pair of auriclelike leaflets;
petiole 3–12 mm; leaf blade suborbicular, cordate, or reniform, (0.7–)1–2.2(–3) × 0.7–2(–2.3)
cm, margin repand or entire, apex rounded; cauline leaves sessile, middle ones (1.5–)3–5(–7) cm;
terminal leaflet orbicular, ovate, or rarely oblong, 1–3 × (0.8–)1–3 cm, with a petiolule
(0.3–)0.5–1.5(–2) cm, base cordate, subtruncate, or obtuse, margin repand or subentire, apex
rounded; lateral leaflets 2–5 pairs, much smaller than terminal one, ovate, suborbicular, or
oblong-ovate; proximal pair of lateral leaflets auriclelike, attached at or just above node, often
giving appearance of amplexicaul leaf base. Racemes ebracteate; fruiting pedicels (0.7–)1–2(–
2.5) cm, slender, divaricate, usually straight. Sepals ovate, 3–3.5 × 1–1.5 mm, margin
membranous, lateral pair subsaccate; petals white, obovate, 7–10 × 3–4 mm, not clawed, apex
rounded to emarginated; filaments slightly spreading, median pairs 4–5 mm, lateral pair 2.5–3.5
mm; anther narrowly oblong, 1–1.5 mm; ovules 14–18 per ovary. Fruit linear, strongly flattened,
(1.5–)2–3(–3.5) cm × 1.5–2 mm; valves smooth, glabrous; styles slender, 1.5–3 mm. Seds
brown, oblong, 2–3 × 1–2 mm, winged all around; wing to 1 mm wide.
Flowering: Apr–Jun.
Habitat: moist places, streamsides.
Elevation: 0–1000 m.
Distribution: China (Anhui, Fujian, Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hunan,
Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Shandong, Sichuan, Zhejiang), Japan, Korea,
Russia (Far East, Siberia).
Cardamine macrocarpa Brandegee, Zoe 5: 233. 1906. TYPE:
Cardamine macrocarpa var. texana Rollins, J. Arnold Arbor. 21: 394. 1940. TYPE: United
States, Texas, Brewster Co., Camp Mountain, Chisos Mountains, Sep 1933, V. L. Cory 7141
(holotype, GH!).
70
Herbs, annual, glabrous or sparsely puberulent. Rhizomes absent; stolons absent. Stems
(1.4–)2–4.5(5.3) dm, erect, or decumbent, 1to or several from base, simple or often branched
above, flexuous or straight, narrowly winged-angled. Basal leaves not rosulate, soon withered;
cauline leaves 3–9, middle ones 5–9-foliolate, 3–8 cm; petiole 1–3 cm, base not auriculate;
terminal leaflet broadly ovate to narrowly lanceolte or rarely oblong, 0.7–2 × 0.2–1 cm, base
cuneate to rounded, margin repand, crenate, or 3-lobed, petiolute 2–8 mm; lateral lobes similar to
terminal, often smaller and with oblique base, upper leaves gradually reduced in size and with
narrower, subsessile leaflets. Racemes ebracteate; rachis slightly to strongly flexuous or
geniculate; fruiting pedicels horizontal to divaricate or ascending, (3–)4–9(–12) mm. Sepals
oblong, 2–3.5 × 0.7–1.2 mm, base of lateral pair not saccate; petals white, linear, 6–8 × 0.7–1
mm; median filament pairs 4–5 mm, lateral pair 3.5–4 mm; anthers oblong, 0.7–1 mm. Fruit
linear, (2.5–)3–4.6 cm × 1.7–2.1 mm; valves glabrous; style 1–3 mm; ovules and seeds 14–22 per
fruit. Seeds dark brown, oblong, 2–2.5 × 0.9–1.2 mm.
Flowering: Mar–Sep.
Habitat: rock crevices and ledges, gravel bars of mountain streams, moist rocky stream banks,
shaded loamy forest floor.
Distribuion: Mexico (Coahuila, Nuevo Leon), United States (Texas/Brewster Co.).
Cardamine macrophylla Willdenow, Sp. Pl. 3: 484. 1800; D. macrophylla (Willdenow) Bunge
ex Maximowicz;
Cardamine macrophylla var. crenata Trautvetter;
Cardamine macrophylla var. dentarifolia J. D. Hooker & T. Anderson;
Cardamine macrophylla var. diplodonta T. Y. Cheo;
Cardamine macrophylla var. foliosa J. D. Hooker & T. Anderson;
Cardamine macrophylla var. lobata J. D. Hooker & T. Anderson;
Cardamine macrophylla var. moupinensis Franchet;
Cardamine polyphylla D. Don 1825, not O. E. Schulz (1903); C. macrophylla subsp. polyphylla
(D. Don) O. E. Schulz; C. macrophylla var. polyphylla (D. Don) T. Y. Cheo & R. C. Fang;
Cardamine macrophylla var. sikkimensis J. D. Hooker & T. Anderson;
Cardamine sachalinensis Miyabe & Miyake;
Cardamine sinomanshurica (Kitagawa) Kitagawa;
Cardamine urbaniana O. E. Schulz, Bot. Jahrb. Syst. 32: 396. 1903. TYPE: China, Sichan,
Augustine Henry 5635 (lectotype, here designated, B!; isolectotypes, GH!, US!).
Dentaria gmelinii Tausch,
Dentaria sinomanshurica Kitagawa,
Dentaria wallichii G. Don,
Dentaria willdenowii Tausch,
Herbs, perennial, Rhizomes creeping, not scaly, slender or stout and with tuberous knots, 2–
10(–30) mm in diam., not stoloniferous Stems (2–)3–9.5(–11.5) dm. stout or slender, erect,
simple or rarely branched above, glabrous, hirsute, or pilose. Rhizomal leaves (4–)10–40(–50)
cm; petiole (1–)3–20(–25) cm; terminal leaflet lanceolate, elliptic, oblong, ovate, or obovate,
(1–)2–15(–25) × (0.5–)1–3.5(–5) cm, sessile or petiolulate, base cuneate, margin serrate, crenate,
dentate, or rarely 3–5-lobed, apex acuminate, acute, or subobtuse; lateral leaflets (1 or)2–6 pairs,
similar to terminal but smaller; cauline leaves 3–12(–18); petiole (1–)2–5(–6.5) cm, not
auriculate at base; terminal leaflet narrowly ovate, lanceolate, elliptic, or oblong, rarely obovate
or lanceolate-linear, (2–)4–12(–20) × 1–4(–5) cm, glabrous or pubescent, with sessile or with a
petiolule to 1 cm, base cuneate, margin ciliolate and crenate, serrate, or serrulate, rarely subentire
or double serrate, apex acute, rarely acuminate; lateral leaflets 2–7(–11) pairs, sessile petiolulate,
71
base cuneate or terminal or all pairs obliquely decurrent, similar to but slightly smaller than
terminal leaflet. Racemes ebracteate, 10–30-flowered; fruiting pedicels ascending or rarely
divaricate, (0.3–)0.8–2.5(–3.1) cm, straight, stout. Sepals oblong, 3.5–6.5(–8) × 1.5–3 mm;
petals purple or lilac, obovate to spatulate, (0.8–)1–1.7 cm × 3.5–8 mm, apex rounded or rarely
subemarginate; median filament pairs 7–9(–11) mm, lateral pair 6–7 mm; anthers oblong, 1–2(–
2.5) mm; ovules 8–12(–16) per ovary. Fruit linear, (2.2–)2.5–6(–7) cm × 1.5–2.5(–3) mm;
gynophore 0.5–3 mm; valves smooth, sparsely pubescent or glabrous; style (1–)2–6.5(–9) mm.
Seeds brown, ovoid to oblong, (1.5–)2–3(–4) × 1–1.7(–2) mm. 2n = 64, 80, 96.
Flowering: (Mar–)Apr–Oct.
Habitat: damp forests, river banks, tundra, rock crevices, meadows, damp woodlands, thickets,
streamsides, valleys, ravines, mountain slopes, amongst boulders.
Elevation: 500–4200 m.
Distribution: Bhutan, China (Anhui, Gansu, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangxi,
Jilin, Liaoning, Nei Mongol, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan,
Zhejiang), India (Lahul, Sikkim), Japan, Kashmir, Kazakstan, Mongolia, Nepal, Pakistan, Russia
(Far East).
Cardamine majovskii Marhold & J. Záborský, Preslia 58: 194. 1986. TYPE:
Cardamine marginata Philippi, Anal. Univ. Chile 27(2): 324. 1865; C. vulgaris var. marginata
(Philippi) O. E. Schulz, Bot. Jarhb. Syst. 545. 1903. TYPE: Chile, Región VIII, Near Chillán,
Manuel Antonio de Solis Obando s.n. (lectotype, here designated, SGO-63903!, fragments,
BAA!). Muñoz (1960) listed another collections that I have not seen, SGO-49376.
Herbs, perennial. Rhizomes much thickened, short, forming a tuberous stem base, not scaly.
Stems solitary, 5−15 cm, erect, simple few branched at middle, glabrous or sparsely pubescent.
Basal leaves and lowermost cauline compound or rarely first leaf simple, 2.5−4 cm, with 3−5foliolate; petiole 1.5−3.5 cm; terminal leaflet 2.5−7 × 1−3 mm, ovate to obovate or suborbicular,
entire, repand or obtusely 3−5-toothed, on a petiolule 1.5−4 mm; lateral leaflets oblanceolate to
linear, 1.5−4 × 0.5−2 mm, sessile or on a petiolule 0.5−2 mm, entire or repand; upper cauline
leaves 3−5-foliolate, smaller than basal; terminal lobe narrowly obovate to oblanceolate, usually
obtusely 3-toothed; lateral lobes linear, sessile. Racemes ebracteate, 3−8-flowered; rachis
flexuous; fruiting pedicels 3−7(−10) mm, divaricate-ascending, slender, straight slightly curved,
forming a distinct angle with fruit. Sepals oblong, 1−1.5 × 0.5−0.7 mm; petals white, narrowly
oblanceolate, 2.5−3.2 × 0.7−1 mm, gradually narrowed to base, apex obtuse; filaments 1.5−2
mm; anthers ovate, 0.2−0.3 mm; ovules 6−10 per ovary. Fruits linear-lanceolate, 1−1.5(−1.7) cm
× 1.8−2.5 mm, attenuate to style; style slender, 1−2 mm. Seeds brown, orbicular to broadly
ovate, 2−2.5 × 1.2−1.5 mm, winged all around; wing 0.1−0.2 mm wide.
Flowering: Sept−Oct.
Habitat: vernal pools.
Elevation: 100−2000 m
Distribution: endemic to Chile (Región VIII).
Specimens examined: CHILE. VIII. Ñuble: 2 km E Chillán on rd to EL Carmen, 36°37′S,
72°5′W, Bliss 512 (CONC), Keeley et al. 25824 (CONC). IX: Malleco, Mininco, [37°47′S,
72
72°28′W], Montero 5254 (CONC, SI), Montero 9416 (CONC). Bío Bío, Antuco, Barros 2790
(SI).
Cardamine marginata resembles the small-sized forms of C. vulgaris, including its type, by
having tuberous stem base, attenuate styles, and fruits forming a distinct angle with the fruiting
pedicels. However, this where the fundamental similarities stop and, indeed, the two species are
very different in flower, infructescence, fruit, and seed morphology. Schulz (1903) reduced C.
marinata to a variety of C. vulgaris and cited a specimen of the latter (Delfin s.n.; SGO) under the
former. The two species are easily distinguished by the presence in C. marginata of much smaller
flowers (sepals 1−1.5 × 0.5−0.7 mm and petals 2.5−3.2 × 0.7−1 mm), flexuous rachis of fruiting
racemes, smaller and wider, linear-lanceolate fruits 1−1.5(−1.7) cm × 1.8−2.5 mm, fewer ovules
(6−10 per ovary), and orbicular to broadly ovate seeds 2−2.5 × 1.2−1.5 mm and winged all
around. By contrast, C. variablis has larger flowers (sepals 2.5−3.5 × 1−1.5 mm and petals
(5−)6−9 × 2−3.5 mm), straight rachis of fruiting racemes, larger and narrower, linear fruits
2−3.5(−4) cm × 1−1.5 mm, more ovules (20−34 per ovary), and ovate, wingless seeds 1.2−1.6 ×
0.8−1.2 mm. Unfortunately, C. marginata is known from only the collections cited above, and a
little can be said about it overall variability.
Cardamine matthioli Moretti, Giorn. Imp. Reale Ist. Lombardo Sci. 8: 623 & Giorn. Imp. Reale
Ist. Lombardo Sci. & Bibliot. Ital. 16: 359. 1847; C. pratensis Linnaeus subsp. matthioli
(Moretti) Nyman, Consp. Fl. Eur. 36. 1878. TYPE: Italy, “le Province di Lombardia,” G.
Moretti s.n. (neotype designated by Marhold & Rayner (1994: 79), FI).
Cardamine maxima (Nuttall) A. Wood, Amer. Bot. & Fl. 38. 1870; Dentaria maxima Nuttall,
Gen. N. Amer. Pl. 2: 66. 1818. TYPE:
Dentaria anomala Eames, Rhodora 5: 217. 1903; Cardamine anomala (Eames) Schumann, Bot.
Jahresber. (Just) 31, pt. 1: 830. 1904. TYPE:
Herbs, perennial, glabrous except for leaflet margin and sometime the raceme rachis.
Rhizomes fleshy, cylindrical, 3–6 mm in diam., distinctly constricted at intervals, not uniform in
diam., slightly fragile, with dentate leaf scars; stolons absent. Stems 0.9–3(–4) dm, erect, simple,
glabrous. Rhizomal leaves 3-foliolate, 7–20 cm; petiole 4–15 cm; terminal leaflet broadly ovate
to oblong, 2–7.5 × 1.2–3.7 cm, with a petiolule 0.2–1(–1.7) cm, base cuneate to obtuse, margin
coarsely dentate to sharply incised or deeply cleft into 2 or 3 lobes, the lobes dentate or incised,
puberulent along margin; lateral leaflets often similar to terminal one in shape, size and margin,
base often oblique, subsessile or petiolulate; cauline leaves 2 or 3, 3-foliolate, alternate or rarely
subopposite; petiole (0.5–)1–4(–6.5) cm, base not auriculate; terminal and lateral leaflets
resembling those of basal leaves, uppermost sometimes much smaller. Racemes ebracteate;
flowering pedicels horizontal to divaricate or deflexed, 0.7–2 cm. Sepals oblong, 5–7 × 2–3 mm,
erect to ascending, base of lateral pair slightly saccate; petals white or pink, oblanceolate, 1–1.7
cm × 3–6 mm, not clawed, apex rounded; median filament pairs 4–8 mm, lateral pair 3–6.5 mm;
anthers linear, 1.7–2.7 mm. Fruit (undeveloped) linear-lanceolate, to 3 cm × 2 mm; valves
glabrous; style 3.5–7 mm; ovules and seeds 10–14 per fruit. Seeds not known. 2n = 120, 124,
132, 138, 156, 161, ca. 208.
Flowering: Apr–Jun.
Habitat: rich woods, shady ravines, ledges, moist alluvial bottoms, steep forested slopes, ledges,
stream banks.
Distribution: Canada (New Burnswick, Ontario, Quebec), United States (Connecticut, Maine,
Massachusetts, Michigan, New Jersey, New York, Vermont).
73
Cardamine mexicana O. E. Schulz, Bot. Jahrb. Syst. 32: 461. 1903. TYPE:
Herbs, annual, hirsute nearly throughout. Stems 1–2 dm, branched about base. Leaves
pinnately compound, 5-foliolate; petioles slender, not auriculate at base; leaflets broadly ovate to
suborbicular or obovate, irregularly crenulate, petiolulate, terminal leaflet largest, 1–2 cm in
diam. Raceme ebracteate, 6–10-flowered, elongated in fruit; fruiting pedicels erect to strongly
ascending, 5–10 mm, slender. Sepals oblong, ca. 2 mm, hirsute; petals white, 3–4 mm; filaments
– mm; ovules – per ovary. Fruit linear, 2–3 cm × 1–1.3 mm, glabrous, erect to ascending; style
thick, < 1 mm. Seeds oblong, wingless.
Flowering Mar–Nov.
Habitat: shady limestone ledges, moist crevices, canyon walls, slopes.
Distribution: Mexico (Coahuila, Nuevo León, San Luis Potosí).
Cardamine micranthera Rollins, Castanea 5: 87. 1940. TYPE: United States, North Carolina,
Stokes Co., Peter’s Creek, near Campbell, 19 May 1940, H. L. Blomquist & G. W. McDowell
11058 (holotype, GH!).
Herbs, perennial, glabrous throughout. Rhizomes very short, 2–3 mm wide, covered with
extensive root system; stolons absent. Stems 0.9–4 dm, erect to ascending, simple, branched
above, glabrous. Rhizomal leaves 3-foliolate or rarely simple, 1–8 cm; petiole 0.5–5 cm;
terminal leaflet or blade os simple leaves orbicular to broadly ovate, 0.5–3 × 0.5–2.5 cm, with a
petiolule 0.4–2 cm, base rounded, margin entire to repand or dentate; lateral leaflets often
minute, subsessile; cauline leaves 5–10, middle ones often simple; petiole 0.3–1.5; blade
rhombic to suborbicular or ovate, 1–3.5 × 0.6–2.2 cm, glabrous, base obtuse to cuneate, margin,
entire, repand, or dentate; upper leaves gradually reduced in size and with shorter petioles.
Racemes ebracteate; fruiting pedicels divaricate, 0.9–1.7 cm. Sepals oblong, 1.5–2.2 × 0.7–1
mm, ascending, base of lateral pair not saccate; petals white, oblanceolate, somewhat spreading,
3.5–5 × 1.2–1.8 mm, not clawed, apex rounded; filaments of median stamens 2.5–3 mm, lateral
filaments 2–2.5 mm; anthers ovate, ca. 0.2 mm. Fruit linear, 0.8–1.6 cm × 0.8–1 mm; valves
glabrous, not torulose; style 1.2–1.8 mm; ovules and seeds 16–22 per fruit. Seeds brown, oblong
to ovate, 0.9–1.2 × 0.6–0.8 mm, wingless.
Flowering: Apr–May.
Habitat: wet grounds along streams, seepage, gravelly sand bars, moist crevices.
Distribution: United States (N North Carolina/Stokes Co.).
Cardamine microphylla J. Adams, Mém. Soc. Imp. Naturalistes Moscou 5: 111. 1817. TYPE:
Cardamine minuta Willdenow ex O. E. Schulz
Herbs, perennial, glabrous or rarely sparsely hirtellous on leaves. Rhizomes slender,
cylindrical, 0.7–1.5 mm in diam.; stolons absent. Stems 0.3–1.5(–2) dm, erect or ascending,
simple, glabrous. Rhizomal leaves pinnately 5–7-foliolate, not fleshy, (1.3–)2.5–6.5 cm; petiole
(0.5–)1.5–5.5 cm; terminal leaflet ovate to elliptic, (2–)4–12(–15) × (1–)2.5–6 mm, glabrous or
rarely pubescent, base cuneate to obtuse, margin entire, apex apiculate, petiolule 1.5–7 mm;
lateral leaflets about as large as or smaller than terminal leaflet, similar in other aspects; cauline
leaves 1–3, 5–7-foliotate, alternate; petiole 2–15 mm, base not auriculate; terminal leaflet ovate
to elliptic, 4–13 × 1–7 mm, subsessile or with a petiolule to 2 mm, base cuneate, margin entire,
apex apicualte; lateral leaflets similar to terminal one. Racemes ebracteate; fruiting pedicels erect
to ascending, 0.7–1.5(–2.5) cm. Sepals ovate to oblong, 3–4 × 1.8–2.5 mm, erect, base of lateral
pair slightly saccate; petals white or rarely lavender, broadly obovate, 7–10 × 3–6 mm, clawed,
apex rounded; median filament pairs 3–4.5 mm, lateral pair 2–3 mm; anthers oblong, 1–1.5 mm.
74
Fruit linear, 2–3 cm × 1.2–1.7 mm; valves glabrous; style 1–2 mm; ovules and seeds 12–16 per
fruit. Seeds brown, oblong, ca. 1.5 × 1 mm. 2n = 28, 42, 64.
Flowering: Jul–Aug.
Habitat: streamsides, sand and cobbles on gravel bars, shale banks, floodplains, alluvial sand
between cobbles, moist moss, turf, seepage areas, meadows, wet grounds.
Elevation: 0–1600 m.
Distribution: Canada (NW Northwestern Territories, N Yukon), Russia (Far East, Siberia),
United States (NE Alaska).
Cardamine microthrix I. Thompson, Muelleria 9: 165. 1996. TYPE: Australia, Victoria, Clarke
Lagoon Wildlife Reserve, NE study area, 28 Oct 1987, A. C. Beauglehole 89710 (holotype,
MEL).
Herbs, annual, glabrous or sparsely pubescent. Stems to 30 cm, often erect, glabrous or
sparsely pubescent, branched at base and above. Basal leaves rosulate, long petiolate, to 8 cm,
3−5-foliolate, leaflets entire, crenate or shallowly lobed; terminaly leaflest broadly ovate, cordate
at bse; lateral leaflets ovate, smaller than terminal, petiolulate; cauline leaves 2 or more, 3−5(−7)foliolate, petiolate, not auriculate at base, leaflets minulately ciliate; terminal leaflet broadly
ovate, often deeply and acutely (3−)5−9-lobed; lateral leaflets ovate, petiolulate, usually 3-lobed.
Racemes ebracteate, few to many flowered; fruiting pedicels suberect, 5−10 mm. Sepals ovate,
ca. 1.5 mm; petals white, cuneate, ca. 3 mm; stamens 6. Frutis linear, erect to suberect, 2−3 cm ×
ca. 1 mm; style 0.5−1 mm. Seeds elliptic, ca. 1.2 mm.
Flowering: Sep−Mar.
Habitat: along banks of rivera, streams, and lagoons and adjacent low-lying areas.
Elevation:
Distribution: Australia (New South Wales, South Australia, Victoria).
Cardamine microzyga O. E. Schulz, Bot. Jahrb. Syst. 32: 545. 1903. TYPE: China, West
Sichuan and Tibetan Frontier, Tachienlu, 9000−13,500 ft, A. E. Pratt 265 (holotype, B!;
isotype, K!).
Cardamine prattii Hemsley & E. H. Wilson, Bull. Misc. Inf. Kew 1906: 153. 1906. TYPE:
China, Sichuan, moist alpine meadows around Tatien Lu, Wilson 3199 (holotype, K!;
isotypes, A, MO!, P!, US!).
Herbs, perennial. sparsely pilose. Rhizomes slender, to 15 cm or longer, 1–1.5 mm in diam.,
without stolons. Stems 10–30(–45) cm, erect, simple, ridged, pilose. Basal leaves rosulate, 2–11
cm; petiole 0.5–5 cm, ciliate; terminal leaflet obovate, 2–8(–13) × 2–7(–10) mm, petiolule 0.5–4
mm, pilose, base cuneate, margin 1–3-toothed on each side, apex acute, submucronate; lateral
leaflets 5–11 pairs, obovate or ovate, pilose, asymmetric, slightly smaller than terminal lobe,
sessile or rarely with a petiolule to 5 mm, base oblique to subcuneate, proximal margin 1- or 2toothed, rarely 1-lobed, distal margin often entire, apex acute; cauline leaves 1–3, (2–)4–10(–25)
cm; petiole 0.5–4(–9) cm, ciliate, not auriculate at base; terminal leaflet and 6–9 pairs of lateral
leaflets similar in morphology to those of basal leaves. Racemes ebracteate, 10–20-flowered.
Fruiting pedicels erect to erect-ascending, (0.7–)1–2 cm, slender, pilose. Sepals oblong or ovate,
2.5–4 × 1–1.7 mm, sparsely pilose or glabrous, margin membranous, base of lateral pair saccate;
petals purple or rarely white, broadly obovate, 6.5–10 × 3–6 mm, cuneate into a clawlike base to
2 mm, apex rounded; median filament pairs 3.5–5 mm, flattened; lateral pair 2.5–3.5 mm;
anthers narrowly oblong, 0.8–1.1 mm; ovules 10–14 per ovary. Fruit linear, 2.5–4 cm × 1.5–2
75
mm; gynophore 0.5–2 mm; valves smooth, glabrous; style 1–2.5 mm. Seeds brown, oblong, 1.8–
2.1 × 1–1.3 mm, wingless.
Flowering: (Apr–)Jun–Sep.
Habitat: moist alpine meadows, streamsides.
Elevation: 2600–4600 m.
Distribution: China (Sichuan, Xizang).
Cardamine moirensis I. Thompson, Muelleria 9: 167. 1996. TYPE: Australia, Victoria, Ulupna
Isalnd, 10 km NW of Stranthmerton in the Murray Valley, eastern end of reserve, 35º51′E,
145º26′S, 20 Sep 1978, T. B. Muir 5965 (holotype, MEL; isotypes, AS, CANB, HO, NSW).
Herbs, annual, glabrous throughout. Stems to 30 cm, slender, erect to ascending, branching
from base and above. Basal leaves rosulate, not fleshy, long petiolate, to 8 cm, 3−9-foliolate;
terminal leaflet ovate, 3-lobed, truncate to cuneate at base; lateral leaflets elliptic to narrowly
ovate, entire or with 1 or 2 lateral teeth; cauline leaves usually 3 or more, petiolate, not auriculate
at base, similar to basal but smaller, leaflets narrowly obovate to filiform and entire in uppermost
leaves. Racemes ebracteate, mostly 2−8-flowered; fruiting pedicels divaricate-ascending, 5−15
mm. Sepals ovate, 1.5−2 mm; petals white, cuneate, 2−3.5 mm; stamens 6. Fruits linear, 1.5−3
cm × ca. 1 mm, suberect; style to 1 mm. Seeds elliptic, 1−1.2 mm.
Flowering: Sep−Oct.
Habitat: low-lying areas near streams and swamps.
Elevation:
Distribution: Australia (New South Wales, Victoria).
Cardamine monteluccii
Cardamine multiflora T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. N. E. Forest. Inst., Harbin
1980(6): 21. 1980. TYPE: China, Sichuan, Paohsin Hsiaen, Mupin, 28 May 1954, Tzupu
Soong 38262 (holotype, NAS!; isotypes, KUN!, PE!).
Herbs, perennial. Rhizomes stout. Stems 35–75 cm, erect, angled, sparsely pilose, lower
nodes sometimes producing stolons to 60 cm, branched above middle. Basal leaves withered by
anthesis; middle cauline leaves pinnatisect; petiole (0.5–)1–3 cm, ciliate, gradually winged from
apex to a much broader, auriculate or amplexicaul base; auricles oblong or ovate, (1.5–)2.5–10(–
13) × 1–3(–5) mm, ciliate; terminal lobe broadly obovate or obovate-oblong, (0.5–)1–4.5(–6) ×
(0.6–)1–2.5(–3) cm, pilose, subsessile or with a petiolule 2–12 mm, base cuneate and often
decurrent with adjacent lateral lobes, margin crenate, apex obtuse to subacute; lateral lobes 1–4
on each side of rachis, similar to terminal lobe but smaller; uppermost leaves and stolon leaves
trifid, with oblanceolate or narrowly obovate terminal lobe decurrent with narrower and smaller
lateral lobes; petiole rarely to 1 cm; auricles rarely to 2 mm or reduced to a tooth. Racemes
ebracteate, many flowered, in panicles; fruiting pedicels divaricate or ascending, 1–2.5 cm,
glabrous, straight. Sepals ovate or oblong, 2–3 × 1–1.5 mm, glabrous, lateral pair subsaccate;
petals purple, spatulate, 6–8 × 2.5–3.5 mm, apex rounded; median filament pairs 3.5–5 mm,
lateral pair 2.5–3.5 mm; anthers oblong, 0.9–1.3 mm; ovules 12–16 per ovary. Fruit linear, 1–2
cm × 1.2–1.5 mm; valves smooth, glabrous; style 1–3 mm. Seeds brown, oblong, 1.5–1.7 × 0.8–1
mm, wingless.
Flowering: May−Jul.
Habitat: mountain slopes, damp places, forests.
Elevation: 2100–3700 m.
76
Distribution: China (Sichuan, Yunnan).
Cardamine multijuga Franchet, Bull. Soc. Bot. France 33: 399. 1886; C. griffithii J. D. Hooker
& Thomson subsp. multijuga (Franchet) O. E. Schulz, Bot. Jahrb. Syst. 32: 506. 1903. TYPE:
China, Yunnan, Mo-soyn, near Lankong, 28 Jun 1884, Delavay 697 (holotype, P!; isotypes,
E!, GH!, P!, US!).
Herbs, perennial, glabrous throughout. Rhizomes creeping, without stolons. Stems (2–)4–
10(–16) dm, erect, simple or branched above, rooting from lowermost nodes, striate angled, 11–
25-leaved. Leaves sessile, cauline; lower and middle ones 2.2–6(–10) cm; terminal leaflet
orbicular or broadly obovate, 3–12 × 3–11 mm, with a petiolule 0.5–6 mm, base subcordate or
cuneate, margin repand and not ciliate, apex obtuse or rounded; lateral leaflets (7 or)8–11(–15)
pairs, slightly smaller than terminal one, ovate or ovate-oblong, base obtuse, margin repand and
not ciliate, apex obtuse or rounded; proximal pair of lateral leaflets auriclelike, attached at or just
above node, often giving appearance of amplexicaul leaf base; uppermost leaves smaller.
Racemes ebracteate; fruiting pedicels divaricate, 1–2 cm, slender, straight. Sepals oblong, 2.5–3
× 1.5–2 mm, spreading; petals purple or lavender, obovate, 7–9 × 3–4 mm, not clawed, apex
rounded; filaments spreading, subequal, 4–5 mm; anthers narrowly oblong, 1.5–1.7 mm; ovules
10–16 per ovary. Fruits (young) glabrous; styles 2–4 mm; stigma entire. Seeds (immature)
wingless.
Flowering Jun–Jul.
Habitat: streamsides, marshy places.
Elevation: 200–2800 m.
Distibution: China (Yunnan).
Cardamine nepalensis Kurosaki & H. Ohba, J. Jap. Bot. 64: 135. 1989. TYPE: Nepal, Janakpur
Zone, Ramechhap district, Serdingma-Dubikharka, 3400–3720 m, 7 Jul 1985, Abies forest, H.
Ohba, T. Kikuchi, M. Wakabayashi, M. Suzuki, N. Kurosaki, K. R. Rajbhandari & S. K. Wu
8570258 (holotype, TI!; isotype, KTM).
Herbs, perennial, shortly pilose or subglabrous. Rhizomes stout. Stems 6–10 dm, erect,
simple, stout, glabrous, 10–18-leaved. Basal leaves not seen; middle cauline leaves petiolate,
minutely auriculate; blade pinnatisect, 4–15 × 1.5–5.5 cm, sparsely pilose; terminal lobe
lanceolate, base cuneate, margin coarsely and irregularly dentate, apex acuminate; lateral lobes
1–3 on each side of midvein, smaller than terminal lobe, decurrent at base, margin denticulate,
apex acute. Racemes ebracteate, 5–25-flowered; flowering pedicels spreading to reflexed;
fruiting pedicels to 1 cm, glabrous. Sepals oblong 4–6 × 1.5–2.5 mm, glabrous or subapically
sparsely pilose, base of lateral pair saccate; petals purple, spatulate to obovate, 1–1.5 cm × 4–7
mm; median filament pairs 7–8 mm, lateral pairs 6–7 mm; anthers oblong, 1–1.5 mm; ovules 10–
14 per ovary. Fruit linear, 2.5–4 cm × 1–2 mm; valves smooth, glabrous; style … mm. Seeds
brown, oblong, ca. 2 × 1 mm, wingless.
Flowering: Jul.
Habitat: Sandy forest floor and streamsides.
Elevation: 2600–3700 m.
Distribution: India (Sikkim), Nepal.
Cardamine niigatensis H. Hara, J. Jap. Bot. 58: 328. 1983. TYPE:
Herbs, perennial. Rhizomes short, creeping, somewhat thickened. Stems (3–)5–15(–20) cm,
erect, glabrous or sparsely pubescent, branched above. Cauline leaves (lower) 1.8–5 cm,
77
trifoliolate or 5-foliolate, glabrous; terminal leaflet orbicular to lanceolate, 0.5–2.3 cm, 0.5–2.5
cm wide, base truncate or obtuse, margin obtusely few toothed, apex rounded; lateral leaflets
elliptic to oblong, 2–10 mm, 1–7 mm wide, base cuneate, margin enite, apex subacute to obtuse;
upper cauline leaves simple or trifoliolate, 2–5 cm; blade or terminal leaflet subsessile or short
petiolate, elliptic, lanceolate, obovate to orbicular, 1.5–4 cm, 0.8–4 cm wide, base obtuse or
subtruncate, entire of crenately 1- or 2-toothed on each side; lateral leaflets obovate to elliptic,
0.5–2 cm, 0.3–0.8 cm wide. Racemes ebracteate; fruiting pedicels 1–1.5 cm, ascending.
Sepals 2–3 mm, 1–1.5 mm wide, glabrous; petals white, 4–7 mm, 2.5–3.5 mm wide; filaments
2.5–3.5 mm. Fruits linear, 2–3.5 cm, ca. 1.2 mm wide, glabrous; style 1–2 mm. Seeds 1–1.2
mm.
Flowering: Apr–May.
Habitat: wet places along streams.
Elevation: 0–1000 m.
Distribution: Japan (Honshu).
Cardamine nipponica Franchet & Savatier, Enum. Pl. Jap. 1: 281. 1879. TYPE: Japan, Kanga,
Monte Hakousan, Ontake, Savatier 2804 (holotype, P!).
Herbs, perennial, glabrous throughout. Rhizomes slender, many branched, with petiolar
remains of previous years. Stems 3–10 cm, erect, 2- or 3-leaved. Basal leaves rosulate, (3 or)5or 7-foliolate; petiole 0.7–2.5 cm; terminal leaflet suborbicular to broadly ovate or elliptic, 1.5–6
× 1–4 mm, base cuneate to obtuse, margin entire, apex acute and submucronate; lateral leaflets (1
or)2–4 pairs, subsessile, obovate, base cuneate, margin entire, slightly smaller than terminal one.
Cauline leaves 3- or 5-foliolate; petiole 4–11 mm, base auriculate to amplexicaul; auricle
lanceolate or toothlike, 0.2–2 × 0.1–0.4 mm, apex acute to subcaudate; terminal leaflet narrowly
oblanceolate to linear, 2–10 × 0.5–1 mm; lateral leaflets 2 pairs, smaller than but similar to
terminal lobe. Racemes ebracteate, 3–6-flowered; rachis flexuous in fruit; fruiting pedicels
divaricate to ascending, 3–8 mm, straight. Sepals oblong, 1.7–2.2 × ca. 0.8 mm; petals white,
spatulate, 5–6 × ca. 1.5 mm, apex rounded; median filament pairs ca. 2.5 mm, lateral pair ca. 1.5
mm; anthers ovate, 0.4–0.5 mm; ovules 10–16 per ovary. Fruit linear, 1.5–3 cm × 0.8–1.2 mm;
valves smooth, glabrous; style 0.8–2 mm. Seeds brown, oblong, 1–1.5 × 0.6–0.9 mm, apically
winged.
Flowering: Jul–Aug.
Habitat: alpine slopes.
Distribution: Japan (Honshu), Taiwan.
Cardamine nuttallii Greene, Bull. Calif. Acad. 2: 389. 1887. TYPE:
Cardamine gemmata Greene, Pittonia 1: 162. 1888; C. californica (Nuttall) Greene var.
gemmata (Greene) O. E. Schulz, Bot. Jahrb. Syst. 32: 387. 1903; C. nuttallii var. gemmata
(Greene) Rollins, Harvard Pap. Bot. 4: 45. 1993; Dentaria gemmata (Greene) T. J. Howell, Fl.
Northw. Amer. 1: 49. 1897. TYPE:
Cardamine tenella var. covilleana O. E. Schulz, Bot. Jahrb. Syst. 32: 391. 1903; C. nuttallii var.
covilleana (O. E. Schulz) Rollins, Harvard Pap. Bot. 4: 45. 1993. TYPE: United States,
California, Siskiyou Mts., 17 Jun 1884, Thomas Howell 30 (lectotype designated by Rollins
(1993b: 285), GH!).
Cardamine tenella var. dissecta O. E. Schulz, Bot. Jahrb. Syst. 32: 391. 1903; C. nuttallii var.
dissecta (O. E. Schulz) Rollins, Harvard Pap. Bot. 4: 45. 1993. TYPE: United States,
78
Washington, Klickitat Co., woods near the Columbia River, 11 Mar-20 Apr 1892, W. N.
Suksdorf 1926 (lectotype designated by Rollins (1993b: 285), GH!).
Cardamine pulcherrima Greene, Erythea 1: 148. 1893; C. nuttalli var. pulcherrima (Greene)
Taylor & Macbryde, Canad. J. Bot. 56: 185. 1978; Dentaria macrocarpa var. pulcherrima
(Greene) B. L. Robinson in A. Gray & S. Watson, Syn. Fl. N. Amer. 1(1); 154. 1895; D.
tenella var. pulcherrima (Greene) Detling, Amer. J. Bot. 23: 573. 1936. TYPE:
Cardamine quercetorum T. J. Howell, Erythea 3: 33. 1895; C. tenella var. querctoum (Howell)
O. E. Schulz, Bot. Jahrb. Syst. 32: 390. 1903; D. quercetorum (T. J. Howell) Greene Pittonia
3: 123. 1896; Dentaria tenella var. quercetorum (T. J. Howell) Detling, Amer. J. Bot. 23: 573.
1936. TYPE:
Dentaria macrocarpa Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 88. 1838, not Cardamine
macrocarpa Brandegee, Zoe 5: 233. 1906. TYPE:
Dentaria tenella Pursh, Fl. Amer. Sept. 2: 439. 1814; Cardamine tenella (Pursh) O. E. Schulz,
Bot. Jahrb. Syst. 32: 389. 1903, not C. tenella E. D. Clarke, Travels 2(1): 117. 1812; C.
pulcherrima var. tenella (Pursh) C. L. Hitchcock, Univ. Wash. Publ. Biol. 17(2); 472. 1964.
TYPE:
Dentaria tenella var. palmata Detling, Amer. J. Bot. 23: 573. 1936. TYPE: United States,
California, Plumas Co., 1877, Mrs. R. M. Austin s.n. (holotype, GH!).
Herbs, perennial, glabrous or sparsely pubescent. Rhizomes slender, producing fleshy, ovoid
to oblong or cylindrical nodal swellings 2–5 mm in diam.; stolons absent. Stems 0.5–2(–3) dm,
erect, simple, glabrous or sparsely pubescent distally. Rhizomal leaves simple or 3(or 5)foliolate, (3–)4–20(–25) cm; petiole (2–)3–18(–21) cm; terminal leaflet or blade of simple leaf
reniform to suborbicular or ovate to oblong, (0.9–)1.3–4(–5.2) × (0.8–)1.2–5(–7) cm, when
compound subsessile or with a petiolule 0.2–3 cm, base cordate to obtuse, margin crenate,
dentate, or 5–7-lobed, with apiculae terminating the teeth or lobes, glabrous; lateral leaflets
(when present) or smaller than terminal leaflet, petiolulate to subsessile, margin same as terminal
leaflet; cauline leaves 1–3, 3(or 5)-foliotate; petiole (0.2–)0.5–2(–3) cm, base not auriculate;
terminal lobe broadly ovate to oblong or linear, (0.5–)1–3.5(–6) cm, petiolulate or sessile and
leaves appearing palmate; lateral leaflets similar to terminal one but smaller, sessile, margin
entire or dentate, rarely lobed. Racemes ebracteate; fruiting pedicels ascending to divaricate, 1–
3.5 cm. Sepals oblong, 3.5–5 × 1.5–2 mm, erect, base of lateral pair saccate; petals purple to pale
pink, rarely white, obovate, 10–15 × 4–7.5 mm, not clawed, apex rounded; median filament pairs
5–8 mm, lateral pair 3.5–5 mm; anthers oblong, 1.5–2 mm. Fruit linear, 2.5–5.6 cm × 2–2.3 mm;
valves glabrous; style 4–8 mm; ovules and seeds 8–16 per fruit. Seeds dark brown, oblong, 2–2.5
× 1.4–1.6 mm.
Flowering Mar–May.
Habitat: open pine forests, damp woods, shaded bottomlands, mossy slopes, streamsides, shaded
and moist hillsides.
Eleavation: 150–1000 m.
Distribution: Canada (British Columbia), United States (California, Oregon, Washington).
Cardamine nymanii Gandoger, Bull. Soc. Bot. France 72: 1043. 1925. TYPE:
Cardamine pratensis Linnaeus var. angustifolia W. J. Hooker, App. Parry J. Sec. Voy. 389.
1825; C. pratensis subsp. angustifolia (W. J. Hooker) O. E. Schulz, Bot. Jahrb. Syst. 32: 529.
1903. TYPE:
Cardamine pratensis subsp. polemonioides Rouy
Herbs perennial, glabrous. Rhizomes caespitose; stolons absent. Stems 0.5–1.6(–3.5) dm,
erect, simple, rarely branched. Leaves thick, with impressed veins; rhizomal leaves with terminal
79
leaflets of the same size and shape as lateral ones, orbicular, broadly ovate to lanceolate, base
rounded to cuneate, mostly entire, petiolulate or sessile, (7 or)9–21-foliolate; cauline leaves 2–
4(–7), pinnatisect or pinnately compound and (7 or)9–21-foliolate, glabrous; petiole base not
auriculate; terminal segments or leaflets of the same shape as lateral ones, narrowly lanceolate to
lanceolate, petiolulate or sessile, base cuneate, margin entire; lateral lobes or leaflets of lower
leaves (4–)7–10 on each side of rachis, fewer upwards, upper leaves with 4 or 5 segments or
leaflets on each side of rachis. Racemes ebracteate; fruiting pedicels erect-ascending, 5–15 mm.
Sepals oblong or ovate, 3.6–4.4 mm, erect, base saccate, green with hyaline margin; petals whitelilac 9–12.3 × 4.8–6.8 mm, clawed, apex rounded or emarginate; median filament pairs 3.5–4.5
mm, lateral 2–3 mm; anthers narrowly oblong 0.9–1.4 mm. Fruit linear, 1–1.8 cm × ca. 1.5 mm;
valves glabrous; style ca. 1 mm, stout; ovules and seeds ca. 16 per fruit. Seeds brown, oblong, ca.
1.5 mm. 2n = 56, 60, 64, 80–100.
Flowering: Jun–Aug.
Habitat: wet meadows, marshes, margins of ponds, along rivers and streams, often along sea
costs, swamps.
Distibution: Canada (Labrador, N Manitoba, Newfoundland, Northwest Territories, Nunavut,
Quebec, Yukon), Europe (Finland, Island, Norway, Sweden), Greenland, Russia (N European
part, Siberia), United States (Alaska). n Eurasia.
Cardamine obliqua A. Richard, Tent. Fl. Abyss. 1: 19. 1847; C. pratensis Linnaeus var.
obliqua (A. Richard), Engler, Hochgebirgsfl. Trop. Afr. 225. 1892. TYPE: Ethiopia,
Begemdir, Schimper II. 989 (holotype, P!; isotypes, B!, BM!, K!, S!, W!).
Cardamine johnstonii Oliver, Trans. Linn. Soc. Bot. 2: 328. 1887. TYPE: Tanzania, Kilimanjaro,
Johnston 21 (holotype, K!; isotype, BM!).
Herbs, perennial, glabrous to densely pubescent. Rhizome creeping, producing few to
numerous shoots. Stems 1–12 dm, decumbent to suberect, many branched baally, persistent,
rooting. Leaves pinnately compound, 4–11-foliolate, 2–17 cm; petioles not auriculate at base;
lateral leaflets elliptic to suborbicular, entrie to dentate or incised, sessile or with petiolules to 2
cm; terminal leaflet larger; uppermost leaves with narrower leaflets. Racemes ebracteate, 5–30flowered; fruiting pedicels ascending 1–2 cm. Sepals 2–4 mm, glabrous, purplish; petals white,
pink or violet, spatulate, 5–9 mm, obtuse at apex; stamens 6; anthers 1–1.5 mm. Fruits linear,
glabrous, 1.4–6 cm × 1–2.1 mm; style 1.8–4 mm. Seeds oblong to suborbicular, 1.2–2.2 × 1–1.5
mm.
Habitat: moist forests, lake shores, streams.
Elevation: 2000–4900 m.
Distribution: Ethiopia, Kenya, Rwanda, Tanzania, Uganda, Zaire.
Notes: the recognition in Rollins (1993b) of some Mexican plants as Cardamine obliqua var.
stylosa Rollins is based on misidentified plants of another species, likely C. ovata.
Cardamine occidentalis (S. Watson ex B. L. Robinson) Howell, Fl. Northw. Amer. 50. 1897;
C. pratensis Linnaeus var. occidentalis S. Watson ex B. L. Robinson in A. Gray, Syn. Fl. N.
Amer. 1: 158. 1895. TYPE: United States,
Cardamine neglecta Greene, Pittonia 3: 154. 1897. TYPE: United States,
Herbs, perennial, glabrous or hirsute. Rhizomes fleshy, ovoid or globose, at base of stem, 3–
10 mm in diam; stolons absent. Stems 1–5 dm, erect to ascending, 1 from base, simple or
branched above, not flexuous, glabrous or pubescent proximally. Basal leaves not rosulate,
pinnately compound, (3 or)5(or 7)-foliolate, 2–10 cm, glabrous; petiole 0.5–6.5 cm; terminal
leaflet orbicular to broadly ovate or subcordate, 0.5–2 × 0.7–2.5 cm, margin entire or repand,
80
base cordate to rounded, petioule 0.3–1.8 mm; lateral leaflets smaller than terminal, ovate, entire,
petiolulate or subsessile; cauline leaves 3–7, middle ones 5–7-foliolate, gradually reduced in size
and becoming 3-folioate upwards; petiole 0.5–3 cm, base not auriculate; terminal leaflet obovate
to oblanceolate, 0.5–2.6 × 0.3–1.3 cm, shallowly toothed or entire to repand, lateral leaflets
smaller. Racemes ebracteate; fruiting pedicels divaricate-ascending, 0.7–1.8 cm. Sepals oblong,
1.7–2 × 1–1.2 mm, base of lateral pair not saccate; petals white, oblanceolate, 4–6 × 1.5–2 mm,
not clawed; median filament pairs 2–2.5 mm, lateral filaments 1–1.5 mm; anthers ovate, 0.3–0.5
mm. Fruit linear, 1.5–3.3 cm × 1.7–2.2 mm; valves glabrous or sparsely pubescent, torulose;
style 0.5–1.5 mm; ovules and seeds 18–40 per fruit. Seeds brown, ovate, 1–1.6 × 1–1.2 mm,
wingless. 2n = 64.
Flowering: Apr–Jul.
Habitat: muddy grounds, lake margins, shallow streams and creeks, meadows.
Elevation: 150–1500 m.
Distribution: Canada (British Columbia), United States (Alaska, N California, Oregon,
Washington).
Cardamine oligosperma Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 85. 1838; C. hirsuta
Linnaeus subsp. oligosperma (Nuttall) O. E. Schulz, Bot. Jahrb. Syst. 32: 468. 1903. TYPE:
United States, “shady woody of the Oregon,” Thomas Nuttall s.n. (holotype, K!; isotypes,
GH!, NY!, PH!).
Cardamine hirsuta var. acuminata Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 85. 1838; C.
acuminata (Nuttall) Rydberg, Bull. Torrey Bot. Club 29: 237. 1902. TYPE: United States,
Oregon, wet places, Thomas Nuttall s.n. (lectotype here designated, BM).
Cardamine hirsuta var. bracteata O. E. Schulz, Bot. Jahrb. Syst. 32: 470. 1903; C. oligosperma
var. bracteata (O. E. Schulz) G. S. Torrey, Rhodora 17: 157. 1915; C. bracteata (O. E.
Schulz) Suksdorf, Rhodora 20: 198. 1918. TYPE: United States, Washington, Klickitat Co.,
low wet places, 8 Apr 1885, W. N. Suksdorf 723 (holotype, B; isotypes, MO!, NY, ORE, US
Cardamine hirsuta var. parviflora Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 85. 1838. TYPE:
United States, Oregon, wet places, Thomas Nuttall s.n. (holotype, BM).
Cardamine oligosperma var. lucens G. S. Torrey, Rhodora 17: 157. 1915; C. lucens (G. S.
Torrey) Suksdorf, Rhodora 20: 198. 1918. TYPE: United States, Washington, Klickitat Co.,
Bingen, 10–12 Apr 1912, W. N. Suksdorf 7452 (holotype, GH!; isotypes, CAS!, MO!, NY!,
ORE, OSC, RM, UTC).
Cardamine unijuga Rydberg, Bull. Torrey Bot. Club 24: 246. 1897; C. hirsuta prol. unijuga
(Rydberg) O. E. Schulz, Bot. Jahrb. Syst. 32: 469. 1903; C. oligosperma var. unijuga
(Rydberg) G. S. Torrey, Rhodora 17: 158. 1915. TYPE: United States, Montana, Gallatin Co.,
Spanish Basin, Madison Range, 6000 ft, 18 Jul 1896, J. H. Foldman 494 (holotype, NY;
isotype, US!).
Herbs, annual or biennial, sparsely hirsute at least proximally. Rhizomes absent; stolons
absent. Stems (0.5–)0.8–3.2(–4.1) dm, erect to ascending, 1 or few from base, simple or
branched above, not flexuous, pubescent throughout or only proximally, very rarely glabrous.
Basal leaves rosulate, persistent by anthesis, pinnately compound, 5–9(–13)-foliolate, 2–8.5(–11)
cm, hirsute or sometimes glabrous; petiole 1–6 cm; terminal leaflet orbicular to ovate or rarely
oblong, 0.4–1.5(–2.3) × 0.3–1(–1.3) cm, margin entire or crenate-dentate to obscurely 3–5-lobed,
base cordate to rounded, petioule 1–7 mm; lateral leaflets smaller than terminal, obovate to
oblanceolate, entire or crentate, petiolulate or subsessile; cauline leaves 3–8, pinnately
compound, similar to basal leaves, gradually reduced in size and number of leaflets upwards;
petiole 0.5–2 cm, base not auriculate. Racemes ebracteate; fruiting pedicels divaricate-ascending,
81
(2–)3–9(–12) mm. Sepals oblong, 1.3–1.8(–2) × 0.5–1 mm, base of lateral pair not saccate;
petals white, narrowly spatulate to oblanceolate, 2.5–3.5 × 0.9–1.5 mm, not clawed; stamens 6,
median filaments 1.7–2.5 mm, lateral filaments 1.2–2 mm; anthers ovate, 0.3–0.5 mm. Fruit
linear, (1.3–)1.6–2.8 cm × 1–1.7 mm; valves glabrous or sparsely pubescent, torulose; style 0.4–
1(–1.5) mm; ovules and seeds 16–36(–42) per fruit. Seeds brown, oblong, 1–1.6 × 0.8–1.2 mm,
wingless. 2n = 16.
Flowering: Mar–Jul.
Habitat: streambanks, shady banks, creek bottoms, lake shores, meadows, moist areas, wooded
slopes.
Elevation: 50–3300 m.
Distribution: Canada (British Columbia), Mexico (Baja California), United States (California,
Colorado, Idaho, Montana, W Nevada, Oregon, Utah, Washington, Wyoming).
Cardamine opizii
Cardamine ovata Bentham, Pl. Hartweg. 158. 1845. TYPE: Colombia, Bogota, Tena, Hartweg
881 (holotype, K!; isotype, G-DEL!).
Cardamine ibaguensis Triana & Planchon, Annal. Sci. Nat. Bot. ser. 4, 17: 60. 1862. TYPE:
Colombia, Antioquia, Manizales, 2700 m, 1851−1857, J. Triana s.n. (lectotype, here
designated, P!).
Cardamine ecuadorensis Hieronymus, Bot. Jahrb. Syst., Beibl. 20: 19. 1895. TYPE: Ecuador,
Loja, Andes, 3000−3200 m, F. C. Lehmann 4826 (holotype, B!; isotype, K!).
Cardamine lehmannii Hieronymus, Bot. Jahrb. Syst., Beibl. 20: 19. 1895; C. ovata subsp.
lehmannii (Hieronymus) O. E. Schulz, Bot. Jahrb. Syst. 32: 412. 1903. TYPE. Colombia,
Cauca, Páramo de Guanacas, 2800−3200 m, F. C. Lehmann 4759 (holotype, B?; isotypes, F!,
GH!, K!, S!).
Cardamine ovata prol. thamnophila O. E. Schulz, Bot. Jahrb. Syst. 32: 413. 1903. TYPE:
Ecuador, “in Silvis Pallatañga Andium Quitensium, Spruce 5555 (holotype, W!; isotype, K!).
Cardamine ovata var. bracteata O. E. Schulz, Bot. Jahrb. Syst. 32: 413. 1903. TYPE: Ecuador.
Pichincha, 4000 m, ?collector 45 (lectotype, here designated, B!).
Cardamine ovata var. eriocarpa O. E. Schulz, Bot. Jahrb. Syst. 32: 413. 1903. TYPE: Ecuador,
Andes, 1857−1859, R. Spruce 5097 (lectotype, here designated, G!; isolectotypes, BM!, NY!,
W!).
Cardamine albertii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 341. 1928. TYPE:
Colombia, Sandtander, edge of Páramo de las Vegas, 3300−3700 m, 20−21 Dec 1926, E. P.
Killip & A. C. Smith 15595 (lectotype, here designated, B; isolectotypes, F!, GH!, K!, NY!).
Cardamine albertii var. minor O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 342. 1928.
TYPE: Colombia, Sandtander, W slopes of Páramo Rico, 3300−3600 m, 15−19 Jan. 1927, E.
P. Killip & A. C. Smith 17766 (holotype, B; isotype, GH!).
Herbs, perennial. Rhizomes often short, neither tuberous nor scaly. Stems (1.5−)2.5−8(−10)
dm, erect or decumbent, simple or branched above, glabrous or sparsely to densely pubescent.
Lower and middle cauline leaves 5−18(−20) cm, (3−)5(−9)-foliolate; petiole 2−9(−13) cm; lateral
leaflets (0.7−)1.5−5(−6) × (0.4−)1−2(−3) cm, lanceolate, oblong, elliptic, ovate, or obovate,
glabrous or sparsely to densely hirsute with trichomes 0.2−0.8 mm, base oblique or cuneate to
subobtuse, margin ciliolate with trichomes to 0.2 mm, serrate or crenate, teeth often mucronate,
apex acute, petiolule absent or to 1.5 cm; terminal leaflets same size or slightly larger than lateral
82
ones, often or a longer a petiolule; uppermost leaves smaller, sometimes trifoliolate or simple
and linear to linear-oblanceolate. Racemes bracteate throughout or basally, rarely only lowermost
flower bracteate; rachis flexuous or straight; fruiting pedicels 1−2.8(−3.5) cm, divaricate,
straight, stout, glabrous or pubescent. Sepals 2−4 × 1−1.5 mm, oblong; petals white or very
rarely pale lavender, obovate to oblanceolate, 5−7 × 2−4 mm, not clawed; filaments 2.5−5 mm;
anthers 0.7−1.4 mm, oblong; ovules 10−20 per ovary. Fruits (2.5−)3.5−5.5(−7) cm ×
1.5−2.5(−3) mm; valves glabrous or sparsely pubescent; style (1−)2−4(−5.5) mm. Seeds 2−3 ×
1−1.5 mm, brown, oblong.
Flowering: Jan−Dec.
Habitat: wet cliff, steep rocky slopes of crystalline schist, cloud forests, open sites, along creeks
and trails in forests, woods, fields, streamsides.
Elevation: 500−3930 m.
Distribution: Bolivia (Cochabamba, La Paz, Larecaja, Santa Cruz), Colombia (Antioquia,
Boyacá, Caldas, Cauca, Cundinamarca, Huila, Magdalena, Meta, Natioquia, Norte de
Dantader, Ocaña, Santander, Tolima, Valle), Costa Rica, Ecuador (Azuay, Carchi,
Chimborazo, Cotopaxi, Imbabura, Loja, Napo, Pichincha, Zamora-Chinchipe), Guatemala,
Honduras, Panama, Peru (Amazonas, Cuzco, Puno) Venezuela (Anzoátegui, Lara, Merída,
Sucre, Tachira, Trujillo).
Specimens examined: BOLIVIA. Cochabamba: Ayopaya, 10 km NW InDep. endencia, Beck &
Seidel 14435 (LPB, MO, SI). La Paz: Franz Tamayo, trail between Pelechuco and Pata, along
Rio Pelechuco, 14°46′S, 69°01′W, Lewis 881704 (G, MO, NY); B. Saaverda, Charazani, 20 km
hacia Apolo, Beck 11379 (MO); Inquisivi, along Río Oscalla between Aquilani-Choquetanga
trail, 10 km N Choquetanga, 16°45′S, 67°17′W, Lewis 39726 (LPB, MO), Lewis 39954 (LPB,
MO); Nor Yungas, near Unduavi, Eyerdam 25238 (F, K); Nor Yungas, between Unduavi and
Chulumani, 12 km E of Unduavi, Croat 51500 (MO); Unduavi, Buchtein 455 (B, BAF, BM, E,
F, G, GH, NY, S); Murillo, Zongo Valley, Solomon 17295 (GH, MO, NY), Solomon et al. 19073
(LPB, MO), Solomon 13100 (GH, MO, NY), Beck 13085 (MO); Murillo, Inca trail, ca. 2 km NE
Challapampa, Ritter & Ritter 1455 (MO). Larecaja: vicinity of Sorata, Challasayo, Mandon 903
(BM, G, GH, K, NY, P, S, W). Santa Cruz: Yungas de San Mateo, Steinbacb 8510 (GH)
COLOMBIA. Antioquia: Mpio de Urrao, Páramo de Frontino, between Llano Grande and
Churrumblúm, 6°28′N, 76°5′W, MacDougal et al. 4379 (GH). Boyacá: Páramo de Belén, Rangel
et al. 4049 (MO); ca. 1 km N of Hacienda Ritacuva, Grubb, Curry & Fernandez-Perez 254 (K,
US); Nevado del Cocuy, Las Lagunillas, Tabloncito, Cuatrecasas 1589 (F). Caldas: above
Salento, Pennell 9344 (GH). Cauca: Cabeceras del río Palo, quebrada de río López, Alto del
Duende, Cuatrecasas 18876 (DS, F, GH, NA); Páramo de Las Paps, El Boquerón, Idrobo, Pinto
& Bischler 3192 (P); Mt. Purace, Pennell & Killip 6675 (GH, NY). Cundinamarca: Macizo de
Bogotá, Cuatrecasas 5384 (F); Muchindote Valley, Alto de Las Cruces, 15 km NE Gachetá,
Grant 9551 (GH, NA); Salto de Tequendamama, Cuatrecasas 190 (F); W slopes of Páramo de
Cruz Verde, Cuatrecasas 362 (F); Río San Francisco, above Bogotá, Pennell 1954 (GH, MO,
NY); Zipaquira, Pennell 2561 (NY). Huila: near mouth of Quebrada la Candela, 25 km SW San
Augustín, 1°45′N, 76°17′W, Fosberg 20079 (GH). Magdalena: Sierra Nevada de Sant Marta,
Los Arroyitos, 73°58′W, 10°56′S, Kirkbride 2366 (NY, US); Quebrada Indiana, Kirkbride 1933
(US). Meta: Macizo de Sumapáz, quebrada El Buque, Díaz et al. 2719 (MO), Díaz et al. 2720
(MO); Sierra de Perijá, E of Manaure, Cuatrecasas & Romero Castaneda 25110 (NY).
Natioquia: 4 kim de Palmitas, Skolnik et al. 19An149 (GH). Norte de Santander: Loso and
vicinity (N Toledo), Killip & Smith 20371 (B, GH, MO, NY, S); between Mutiscua and
83
Pamplona, Killip & Smith 19704 (GH) Ocaña: New Grenada, Schlim 303 (G, P). Santander:
Páramo de romeral, Killip & Smith 18566 (GH, NY)12 km N of Vélez, Fassett 25303 (GH).
Tolima: Rosalito, near Páramo de Ruis, Pennell 3118 (MO, NY). Valle: Hoya del río Cali,
confluence of río Pichindecito with el Pichindé, Cuatrecasas 18766 (US). ECUADOR. Azuay:
Giron, Parroquia Asunción, Bosque seco Montano Bajo, 79°15′W, 03°09′S, Cerón 14463 (MO);
Nudo de Portete, pass between headwaters of ríos Tarquic and Giron, Camp E-2170 (GH, NY).
Carchi: Mira Canton, 30 Km al Oeste de San Isidro, 78°05′W, 0°35′S, Gudiño 441 (MO,
QCNE); Espejo, Camino Libertad-Morán, Nacimiento del Río Morán, 78°02′W, 0°45′S, Palacios
11768 (MO, QCNE); Espejo, Reserva Ecológica El Angel, Palacios 11554 (MO,QCNE); Valle
de Maldonado, km 53 on rd Tulcán-Maldonado, 0°50′N, 78°3′W, Holm-Nielsen et al. 5809
(NY). Chimborazo: Camino Pusucucho al Placer, Acosta Solís 7250 (F), Acosta Solís 7274 (F).
Cotopaxi: Salcedo, Parque Nacional Llanganates, E of Salcedo, valley of Rio Anatenorio, km
56, 0°59′0″S, 78°17′2″W, Neil, Vargas, Freire & Narvaes 11903 (MO, NY); Cantón Sigchos,
Orillas del Río Los Illinizas, dentro del Bosque, 78°47′W, 00°34′S, Ramos et al. 6007 (MO).
Imbabura: Pimampiro Canton, Ibarra-Mariano Acosta, Loma Yanalpacunga, 78°01′W, 0°18′S,
Palacios & Tipaz 9903 (MO, QCNE); between Las Toldadas and Río Arturo, 2 mile NE
Cayambe Peak, Wiggins 10512 (DS, GH); shore of Laguna de San Marcos, Wiggins 10523 (DS,
GH, NY); Río San Pedro, below mouth of Río Clavadero, Wiggins 10455 (DS, GH); Lago San
Marcos, Cazalet & Pennington 5311 (B, K, NY, UC, US). Loja: Parque Nac. de Podocarpus,
along trail to Lag. de Compadre, 79°08′W, 04°03′S, Laegaard 101900B (AAU); trails ca. 5 km
ENE of San Pedro de Vilcabamba, from El Bosque to Quebrada Romerillos and Banderilla,
79°10′W, 04°14′S, Øllgaard & Navarrete 105966 (AAU). Napo: Cantón Quijos, 4 km al N de
Papallacta, 78°09′W, 00°20′S, Yánez & Maza 906 (MO); Quito-Baeza road, 5 km NW of
Laguna de Papallacta (Páramo de Guamani), 78°07′W, 0°19′S, Øllgaard & Balslev 8065 (AAU,
MO, NY); Volcán Cayambe, 77°59′W, 0°03′S, Øllgaard et al. 34217 (AAU, MO, NY, QCA);
Laguna San Marcos, 77°58′W, 0°07′S, Øllgaard et al. 34104 (AAU, MO, NY, QCA); S Cerro
Sumaco, 0°36′S, 77°39′W, Løjtnant & Molau 13095 (NY). Pichincha: 2 km S Atahualpa,
Asplund 20302 (NY, P, S); between Nono and Nanegal, Asplund 17324 (B, K, NY, S);
Cayambe-Olmedo-Laguna San Marcos, 77°57′W, 0°07′S, Jørgensen 61958 (AAU, MO, NY,
QCA), Cuamacás & Gudiño 467 (MO), Cuamacás & Gudiño 546 (MO); Parroquia Calacali,
Reserva Geobotánica Pululahua, 78°30′W, 0°05′S, Cerón & Regalado 6156 (MO); Tbacundo,
Lagunas Mojanda, Palacios, Tipas & Aulestia 10209 (MO, QCNE); Campamento Sigsal between
San Juan and Chiriboga, Asplund 16230 (NY). Zamora-Chinchipe: Km 17 on Loja-Zamora rd,
79°08′W 03°59′S, Holm-Nielsen et al. 3546 (AAU, MO, NY). PERU. Amazonas: Bagua Distr.,
Aramango, Catarata Numparket Claro, 78°19′W, 05°29′S, Vásquez, Rojas & Campos 27262
(MO); Bagua, Cordiller Colán SE La Peca, Barbour 3708 (F, MO, NY); Bagua, ca. 25 km E La
Peca, Barbour 2878 (MO). Cuzco: Urubamba, 101 km from Cuzco, between Pampacahua and
Cedrobamba, Quebrada and Rio Pachachoca, Núñez et al. 11090 (MO); Urubamba, Santuario
Histórico Machu Pichu, 72°07′W, 13°18′S, Núñez & Pyrke 9211 (MO); Ollantaytambo,
Huaytampo, 72°21′W, 13°10′S, Calatayud et al. 959 (MO); La Convención, Santa Teresa,
Choqekiraw, 13°23′S, 72°52′W, Valenzuela et al. 3505 (MO). Puno: Prov Sandia, Limbani,
Vargas 9652 (K, NA, UC). Dep. t.??: Tambo de Vaca, Macbride 4462 (F, GH, NY); Chaglla,
Macbride 3637 (F, NY). VENEZUELA. Anzoátegui: along Río Zumbador, near base of Piedra
Blanca, NE Bergantín, Steyermark 61316 (F). Lara: Dtto. Moran, trail from Humocaro to
84
Buenas Aires (Caserio), below Páramo Los Rosas, 70°05′W, 09°40′N, Liesner et al. 8161 (MO);
between Buenos Aires and Páramo de las Rosas, Steyermark 55450 (F, GH); Parque Nac. Dinira,
Páramo de Jabón, 70°06′W 09°34′N, Riina, Meier & Duno 913 (MO, VEN). Merida: from cabin
at La Escalera to Puent de La Escalera, Luteyn, Lebrón-Luteyn & Ruíz-Terán 6227 (MO, NY);
Del Puente de la Quebrada el Plan hacia Aricagua, Los Magos, Bernardi 6193 (MO); between
Los Corales and Las Cuadras, Steyermark 55754 (F, K, S); Jiménez, 10-15 km SSE Sanare,
between Encrucijaa and rd of Parque Nacional Yacambú de El Blanquito, Steyermark et al.
10351 (W). Sucre: bse of Cerro de Diablo, W extension of S peak of Cerro Turumuquire,
Steyermark 62743 (F, GH). Tachira: along Quebrada Agua Azul, S El Reposo, 14 km SE
Delicias, 72°24′W, 7°31′N, Steyermark & Liesner 118383 (GH, MO, NY); Quebrada La Lejia,
15 km SE Delicias, Steyermark & Liesner 118544 (GH, MO); Chorrejon de la Mota de la Peña
de Ventana, Steyermark & Dunsterville 100800 (US). Trujillo: Parque Nacional Dinira, Mun.
Carache, paramo de Jabon to Mesa Arriba, 70°8′W, 9°34′N, Meier, Riina & Duno 6271 (MO,
VEN); between Mesa Arriba and Páramo de Las Rosas, Duno & Riina 1402 (MO, VEN);
Boconó, Cerro Guaramacál, Stergios, Aymard & Smith 4745 (MO); Boconó, Paramo de
Guaramacal, 25 km SE Boconó, 70°10′W, 9°13′N, Aymard, Ortega & Moran 2961 (MO, NY);
12 km E-SE Boconó, 70°9′W, 9°12′N, Liesner et al. 12936 (MO); Boconó, 20-22 km on rd to
Guaramacal, 70°13′W, 9°13′N, Luteyn & Cotton 9745 (GH, MO, NY); Parque Nacional
Guaramacal, rd from Boconó to Guaramacal, SE Boconó, 70°12′W, 9°13′N, Dorr & Barnett
8253 (MO, NY, US).
As in several other South American species of Cardamine, C. ovata is highly variable in leaflet
shape, size, margin, and pubescence (see description above), and the synonymy above reflects
taxa based on some of that variation. In some forms, the leaflets are completely glabrous,
whereas in others (e.g., Lehmann 4826, the type collection of C. ecuadorensis) they are very
densely pubescent. In rare occasions when the plants are pubescent, (but not in Lehmann 4826),
the fruits too are pubescent, as in the types of the Ecuadorian infraspecific taxa thamnophila and
eriocapa. Fruit pubescence is extremely rare in Cardamine as a whole, but these Ecuadorian
plants are indistinguishable in other characters from the remainder of C. ovata.
***********************
These are annotated by Schulz as:
C. jamesonii prol. decora O.E. Schulz: Colombia, Cauca, Paramo de Guanacas, Lehmann 2116
(G).
C. johnstonii: Colombia, Pasto, Lehmann 525a (G)
C. ovata subsp. lehmanniana var. bracteata O. E. Schulz: Colombia, Cauca, Paramo del Ruiz,
Lehmann 3108 (G).
***********************
Cardamine pachystigma (S. Watson) Rollins, Harvard Pap. Bot. 4: 45. 1993; Dentaria
californica Nuttall var. pachystigma S. Watson, Proc. Amer. Acad. Arts 14: 289. 1879;
Cardamine californica (Nuttall) Greene var. pachystigma (S. Watson) O. E. Schulz, Bot.
Jahrb. Syst. 32: 388. 1903; D. pachystigma (S. Watson) S. Watson in A. Gray & S. Watson,
Sy. Fl. N. Amer. 1(1): 155. 1895. TYPE:
Dentaria corymbosa Jepson, Man. Fl. Pl. Calif. 426. 1925, non Matsumura,... 1899. TYPE:
85
Dentaria corymbosa var. grata Jepson, Fl. Calif. 2: 57. 1936. TYPE:
Herbs, perennial, glabrous. Rhizomes fleshy, ovoid to oblong, (3–)4–18 mm in diam., deeply
underground; stolons absent. Stems 1–3 dm, erect, simple, glabrous. Rhizomal leaves simple,
1.2–2.3 cm; petiole 6.7–19.5 cm; blade orbicular to reniform or cordate to broadly ovate, 2.7–5.5
× 3–6.8 cm, base cordate to subtruncate, margin with obtuse teeth terminated in apiculae; cauline
leaves 2–5, crowded above middle of stem, simple; petiole 0.3–2.6 cm, base not auriculate; blade
similar in shape and margin to that of basal leaves, 1.8–5.5 × 1.2–4.7 cm, glabrous, base cuneate
or obtuse to subcordate, marginal teeth apiculate. Racemes ebracteate; fruiting pedicels
divaricate to ascending, 0.7–2.4 cm. Sepals oblong, 4–7 × 2–2.5 mm, erect, base of lateral pair
saccate; petals pink or purple or rarely white, obovate to oblanceolate, 14–18 × 5–7 mm, claws to
9 mm, apex rounded; median filament pairs 7–8 mm, lateral pair 5.5–6.5 mm; anthers oblong, ca.
1.5 mm. Fruit linear, 3.2–5.4 cm × 2.2–4 mm; valves glabrous; style 4–7 mm; ovules and seeds
10–14 per fruit. Seeds brown, broadly ovate, 2–2.8 × 1.8–2.5 mm.
Flowring: Mar–May.
Habitat: forests, lava slides, talus and cliffs.
Elevation: 700–2900 m.
Distribution: United States (E and NC California/Plumas, Tehama, and Tulare counties).
Cardamine papillata I. Thompson, Muelleria 9: 161. 1996. TYPE: Australia, Victoria,
Maramingo Creek area, ca. 4 km NE of Genoa P.O., 26 Dec 1969, A. C. Beauglehole 32819
(holotype, MEL).
Herbs, annual, glabrous or papillate. Stems to 25 cm, erect to ascending, slender, glabrous or
minutely papillate, often branched below and above. Basal leaves rosulate, long petiolate, simple
or 3−5-foliolate, to 7 cm; terminal leaflet orbicular, slightly cuneate or often shallowly cordate at
base, entire; lateral leaflets much smaller than terminal, short petiolulate or sessile, orbicular;
cauline leaves 0−3, sessile, not auriculate at base, lowermost similar to basal, simple or 3−5foliolate, much reduced in size upwards, pinnatifid or pinnate. Racemes ebracteate, 1−10(−15)flowered; fruiting pedicels divaricate-ascending, 5−15 mm, glabrous or minutely papillate.
Sepals ovate, 1.5−2.5 mm; petals white or pink outside, spatulate, cuneate, 3−4.5 mm; stamens
6. Fruits linear, erect to suberect, 2−3.5 cm × ca. 1 mm; style to 1 cm. Seeds elliptic, 1−1.2 mm.
Flowering: Aug−Feb.
Habitat: hilly, forested areas at lower altitudes, moist shaded areas often in rocky siltes near
streams.
Elevation: 0−450 m.
Distribution: Australia (New South Wales, South Australia, Tasmania, Victoria).
Cardamine papuana (Laut.) O. E. Schulz, Bot. Jahrb. Syst. 55: 271. 1903; C. africana Linnaeus
subsp. borbonica (Persoon) O. E. Schulz var. papuana Laut. in K. Sch. & Laut., Fl. Deut.
Schutzgeb. Südsee, Nachtr. 271. 1905. TYPE:
Herbs, perennial, glabrous or rarely pubescent. Rhizomes short, not fleshy. Stems 1.5–5 dm,
erect to ascending, simple or branched above. Basal leaves and lowermost cauline trifoliolate or
unifoliolate, to 12 cm; petiole to 9 cm; cauline leaves few, petiole, 2–6 cm, minutely auriculate or
not; terminal leaflet lanceolate, 3–12 × 1–4 cm, cuneate to rounded or rarely truncate at base,
serrate-dentate, acute to acuminate at apex, with a petiolule 0.5–2 cm; lateral leaflets ovate, 0.5–3
× 0.2–2 cm, serrate to dentate, on a petiolule 1–4 mm. Racemes ebracteate, 5–15-flowered,
elongated or not in fruit; fruiting pedicels ascending to divaricate. Sepals oblong, 2.5–4 mm;
petals white, broadly spatulated, 8–12 × 3–5 mm, usually notched; stamens 6, anthers ca. 0.8
86
mm; ovules – per ovary. Fruits linear, 2–4 × 1.5–2 mm; style 0.5–1.7 mm. Seeds oblong, 2.7–2
× 1–1.3 mm, smooth.
Habitat: rocky banks of forest streams, wet cliffs.
Elevation: 500–3100 m.
Distribution: New Guinea.
Cardamine parviflora Linnaeus, Syst. Nat., ed. 10, 2: 1131. 1759. TYPE: Herb. Linn. 835.10
(lectotype designated by Jonsell (1982: 43), LINN).
Cardamine arenicola Britton, Bull. Torrey Bot. Club 19: 220. 1892; C. parviflora var. arenicola
(Britton) O. E. Schulz, Bot. Jahrb. Syst. 32: 485. 1903. TYPE:
Cardamine brachycarpa Franchet, Bull. Soc. Bot. France 26: 82. 1879, not Opiz, Natural. 11:
411. 1826; C. koshiensis Koidzumi, Fl. Symb. Orient.-Asiat. 43. 1930. TYPE: Japan, Prov.
Etchigo, near Niigata, R. P. Faurie 23 (holotype, P!; isotype, P!).
Cardamine flexuosa Withering var. gracilis O. E. Schulz, Bot. Jahrb. Syst. 32: 481. 1903. TYPE:
Cardamine manshurica (Komarov) Nakai, Tyosen-Syobubutu 113. 1914; C. parviflora Linnaeus
var. manshurica Komarov, Trudy Imp. S.-Peterburgsk. Bot. Sada 22: 370. 1903. TYPE:
Herbs, annual, slender, glabrous or sparsely to densely pubescent throughout. Rhizomes
absent; stolons absent. Stems (0.5–)1–3(–4) dm, erect, 1 to several from base, often branched
above, somewhat flexuous. Basal leaves usually not rosulate, often withered by anthesis,
pinnately (5–)7–13(–17)-foliolate, (2–)4–10 cm; petiole 0.5–2.5(–4.5) cm; terminal leaflet linear
to oblong or oblanceolate to obovate or suborbicular, (1–)3–10 × 1–7 mm, base cuneate, margin
entire, or 3(–5)-toothed or –lobed, sessile or petiolule to 5 mm; lateral leaflets similar to terminal
or sometimes smaller; cauline leaves 5–10(–14), (5–)9–15(–17)-foliolate; petiole 3–10 mm, base
not auriculate; terminal leaflet filiform, linear, or narrowly oblong, 3–10(–16) × 0.3–3 mm,
sessile, entire or rarely 1–3-toothed; lateral leaflets similar to terminal or slightly smaller.
Racemes ebracteate; fruiting pedicels divaricate or ascending, 4–10 mm. Sepals oblong, 1–1.5(–
2) × 0.3–0.5 mm, membranous at margin; petals white, oblanceolate, (1.5–)1.8–2.5(–3) × 0.4–
0.8(–1) mm; stamens 6; filaments 1.4–2.5 mm; anthers ovate, 0.2–0.4 mm. Fruit linear, (0.5–)1–
2(–2.5) cm × 0.6–0.9 mm; valves glabrous, torulose; style 0.3–0.7(–1) mm; ovules and seeds 20–
50 per fruit. Seeds pale brown, oblong-ovate, 0.6–0.9 × 0.4–0.6 mm, narrowly margined or not.
2n = 16.
Flowering: Mar–Jun.
Habitat: roadsides, river banks, rocky crests and outcrops, crevices of granitic bedrock, dry
woods, glades, fallow fields, disturbed grounds, limestone barrens, marsh and swamp
margins, floodplains, waste grounds, slopes, ledges and cliffs, meadows.
Elevation: 0–1500 m.
Distribution: Algeria, Austria, Azerbaijan, Bulgaria, Canada (Alberta, British Columbia,
Manitoba, New Brunswick, Newfoundland, Northwest Territories, Ontario, Quebec,
Saskatchewan), China (Anhui, Guangxi, Hebei, Heilongjiang, Hunan, Hubei, Jiangsu,
Liaoning, Nei Mongol, Shaanxi, Shandong, Shanxi, Xinjiang, Zhejiang), Czech Republic,
Finland, France, Georgia, Germany, Hungary, Iran, Italy, Japan, Kazakhsta, Korea, Mongolia,
Poland, Portugal, Romania, Russia (European Part, Siberia), Spain, Sweden, Taiwan, Ukraine,
United States (Alabama, Arkansas, Connecticut, Delaware, District of Columbia, Florida,
Georgia, Idaho, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland,
Massachussets, Michigan, Minnesota, Missouri, New Hampshire, New Jersey, New York,
North Carolina, Ohio, Oklahoma, Oregon, Pennsylvania, Rhode Island, South Carolina,
Tennessee, Texas, Vermont, Virgina, West Virgina, Wisconsin).
87
Cardamine pattersonii L. F. Henderson, Rhodora 32: 25. 1930. TYPE:
Herbs, short-lived perennial or sometimes annual, glabrous throughout. Rhizomes slender,
cylindrical, 0.5–1.5 mm in diam.; stolons absent. Stems 0.6–3 dm, erect, single, simple or
branched near or at base, not flexuous, glabrous. Basal leaves rosulate or not, 3–5-foliolate, 1–6
cm; petiole 0.4–2.5 cm; terminal leaflet obovate to orbicular or subcordate, 3–15(–20) × 2.5–
16(–18) mm, base obtuse to rounded or cordate, margin entire or dentate to slightly sinuately
lobed, petiolule 1–4 mm; lateral leaflets similar to terminal but considerably smaller; cauline
leaves 2–4 below the raceme, 3–5-foliolate, similar to basal leaves, gradually reduced in size as
bracts, uppermost ones subtending pedicels of uppermost flowers almost always simple; leaflets
or simple bracts linear to linear-oblanceolate, 2–8 mm. Raceme bracteate throughout; fruiting
pedicels divaricate-ascending, (1–)1.5–3(–4.5) cm. Sepals oblong, (–2–3 × 0.7–1 mm, lateral
pair not saccate; petals purple or pink, obovate, 6–9 × 3–4 mm, not clawed; median filaments 3–
3.5 mm, lateral filaments 2–2.5 mm; anthers oblong, 0.7–1 mm. Fruit linear, 2–3 cm × 1–1.5
mm; valves glabrous, torulose; style 2–4 mm; ovules and seeds 14–20 per fruit. Seeds brown,
oblong to ovate, 1.7–2.2 × 1–1.5 mm, distally winged.
Flowering: May–Jun.
Habitat: moist mossy cliffs, rocky slopes, mossy banks.
Elevation: 800–900 m.
Distribution: United States (NW Oregon/Clastop Co.).
Cardamine paucifolia Handel-Mazzetti, Symb. Sin. 7: 359. 1931. TYPE: China, Yunnan, near
Yungbei, between Dawan and Gwangyilang, 2400−2600 m, 3 Jul 1914, Handel-Mazzetti
3432 (holotype, WU!; isotype, W!).
Cardamine yunnanensis Franchet var. obtusata C. Y. Wu ex T. Y. Cheo & R. C. Fang, Bull. Bot.
Lab. North-East. Forest. Inst., Harbin (1980)6: 20. 1980. TYPE: China, Yunnan, Kang-pu,
Wei-si Hsien, C. W. Wang 64172 (holotype, KUN!; isotypes, A!, NAS!).
Herbs, perennial, sparsely pilose or glabrescent. Rhizomes slender. Stems 15–40 cm, simple,
slightly angled. Basal leaves often withered by flowering, trifoliolate; petiole 1.5–4.5 cm; leaflets
similar to those of cauline leaves, smaller; cauline leaves 1–4, trifoliolate; petiole 1.5–6 cm, not
auriculate at base; terminal leaflet lanceolate, ovate, or broadly elliptic, 3–6.5 × 1–3.5 cm,
sparsely pilose or glabrous, with a petiolule 5–12 mm, base obtuse or cuneate, margin ciliate and
dentate or crenate, apex acute or acuminate; lateral leaflets 2, shortly petiolulate or sessile and
base decurrent to rachis, nearly as large as terminal one, base oblique; uppermost leaf often with
a sessile terminal leaflet decurrent basally with lateral leaflets. Racemes ebracteate; fruiting
pedicels ascending or divaricate, 7–11 mm, straight, slender. Sepals oblong, 3.5–4.5 × 1.2–1.5
mm, base not saccate; petals white, narrowly obovate, 6–8.5 × 1.5–3.5 mm, not clawed; median
filament pairs 3.5–4 mm, lateral pair ca. 3 mm; anthers oblong, 1.1–1.6 mm; ovules 10–16 per
ovary. Fruit linear, 3–5 cm × 1–1.5 mm; valves smooth, glabrous; style 1.5–2.5 mm. Seeds
brown, narrowly oblong, 1.5–2 × 0.6–0.8 mm, apically winged.
Flowering: Mar−Jul.
Habitat: ravines, streamsides.
Elevation: 1500–2600 m.
Distribution: China (Yunnan).
Cardamine paucijuga Turczaninow, Bull. Soc. Imp. Naturalistes Moscow 27: 295. 1854.
TYPE: SW Australia, J. Drummond 5: 131 (holotype, KW; isotypes, G, K).
88
Cardamine inermedia subsp. antipodum O. E. Schulz, Bot. Jahrb. Syst. 32: 486. 1903. TYPE:
Australia,
Herbs, annual, glabrous or sometimes pubescent. Stems 1–4 dm, slender, erect. Basal leaves
and cauline pinnately compound, with 1–4 leaflet pairs; leaflests narrowly obovate to elliptic, not
ciliate, terminal largest, not lobed, base cordate; cauline leaves 2 or more, petiolate, not
auriculate at base, leaflets elliptic to narrowly obovate, entire. Racemes ebracteate, often more
than 8-flowered; fruiting pedicels spreading to subhorizontal, slender, 3–15 mm. Sepals oblong,
1.5–2 mm; petals white or rarely pink, 3–6 mm; stamens 6; ovules – per ovary. Fruits linear, 1–3
cm × 0.7–1 mm, erect; style slender, to 2 mm. Seeds ovoid to oblong, 1–1.5 mm, flattened, nery
narrowly winged.
Flowering: Oct–Nov.
Habitat: dry to moist areas in rich grounds.
Distribution: Australia (all states except Northern Territories).
Cardamine pectinata
Cardamine pedata
Cardamine penduliflora O. E. Schulz, Bot. Jahrb. Syst. 32: 538. 1903. TYPE:
Cardamine rariflora M. E. Peck, Torreya 32: 150. 1932. TYPE:
Herbs, perennial, glabrous throughout. Rhizomes fleshy, tuberous, 4–9(–11) mm in diam.;
stolons absent. Stems 2–6(–7.5) dm, erect or decumbent at base, simple, glabrous. Rhizomal
leaves 5–13-foliolate, (4–)10–18(–25) cm; petiole (3–)5–12(–17) cm; terminal leaflet oblong to
elliptic or ovate, (0.4–)0.7–1.7(–2) cm, with a petiolule 4–7 mm, base cuneate or obtuse, margin
entire or obscurely 3-lobed, glabrous; lateral leaflets about as large as or smaller than terminal
leaflet, subsessile, margin same as terminal leaflet; cauline leaves 2–6, 5–11-foliotate; petiole 3–
10 cm, base not auriculate; terminal lobe narrowly ovate or oblong to oblanceolate, 1.5–3.5 ×
0.2–1.5 cm, glabrous, entire or toothed to 3-lobed, petiolule 0.5–1.5 cm; lateral leaflets similar to
terminal one but smaller, sessile. Racemes ebracteate; fruiting pedicels ascending to divaricate,
(1–)2–4(–6) cm. Sepals oblong to ovate, 3.5–5 × 1.8–2.5 mm, erect, base of lateral pair saccate;
petals white, obovate, 12–16 × 6–8 mm, not clawed, apex rounded or subemarginate; median
filament pairs 6–7 mm, lateral pair 4–5 mm; anthers oblong, 1.5–1.8 mm. Fruit linear, 2.5–4.5
cm × 1.4–2 mm; valves glabrous; style 4–6 mm; ovules and seeds 12–24 per fruit. Seeds brown,
oblong, 1.8–2 × 1–1.5 mm.
Flowering: Mar–May.
Habitat: shallow pools, wet grounds, marshes, meadows, creeks, channels, swampy woods.
Elevation: 50–150 m.
Distribution: United States (W Oregon).
Cardamine pensylvanica Muhlenberg ex Willdenow, Sp. Pl. 3: 486. 1800; C. flexuosa
Withering subsp. pensylvanica (Muhlenberg ex Willdenow) O. E. Schulz in Urban, Symb.
Antill. 3: 521. 1903; C. hirsuta Linnaeus var. pensylvanica (Muhlenberg ex Willdenow) P. W.
Graff, Castanea 10: 95. 1945; Dracamine pensylvanica (Muhlenberg ex Willdenow)
Nieuwland, Amer. Midl. Naturalist 4: 40. 1915. TYPE: United States, Pennsylvania,
Muhlenberg s.n. (holotype, B-WILLD).
Cardamine oregana Piper, Proc. Biol. Soc. Wash. 37: 92. 1924; C. breweri var. oregana (Piper)
Detling, Amer. J. Bot. 24: 73. 1937. TYPE: United States, Oregon, near Little Meadows of the
89
Deschutes River, 1500 m, 25 Aug 1896, J. B. Leiberg 2614 (holotype, US!; isotypes, GH!,
NY!).
Cardamine multifolia Rydberg, Bull. Torrey Bot. Club 29: 238. 1902. TYPE: United States,
Utah, Wasatch Mts., 26 Jun 1869, S. Watson 80 (lectotype designated by Holmgren (2004b:
245), NY!).
Cardamine pensylvanica var. brittoniana Farwell, As Gray Bull. 2: 46. 1894. TYPE: United
States,
Cardamine rotundifolia Michaux var. diversifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 427. 1903.
TYPE: United State,
Herbs, annual or biennial, glabrous above, sparsely hirsute near base. Rhizomes absent;
stolons absent. Stems (0.5–)1.5–5.5(–7) dm, erect, single, simple or branched above, not
flexuous, sparsely hirsute hirsute near base, glabrous above. Basal leaves soon withered, not
rosulate, 4–15 cm, similar to lowermost cauline leaves; cauline leaves (3–)5–20(–35), pinnately
(5–)7–13(–19)-foliolate, sometimes appearing pinnatisect and with lobe number similar to that of
leaflets, middle and lower ones 2–11 cm; petiole (0.4–)1–3.5(–4.5) cm, often sparsely hirsute,
base not auriculate; terminal leaflet suborbicular or obovate to oblanceolate or elliptic, 1.3–3(–4)
× 0.6–2.5 cm, base often cuneate, margin entire to repand or obscurely 3–5-lobed, subsessile or
petioule to 10 mm; lateral leaflets smaller than terminal, entire or crenate, short petiolulate or
sessile and decurrent on rachis; uppermost leaves narrower and with fewer lobes or leaflets.
Racemes ebracteate; fruiting pedicels divaricate-ascending, (3–)4–10(–13) mm. Sepals oblong,
(1–)1.3–2.3 × 0.5–1 mm; petals white, narrowly spatulate to oblanceolate, 2–3.5(–4) × 0.8–1.5
mm, not clawed; stamens 6, median filaments 1.5–2.5 mm, lateral filaments 1–2 mm; anthers
ovate, 0.2–0.3 mm. Fruit linear, (1.4–)1.7–2.7(–3.2) cm × 0.8–1.1 mm; valves glabrous,
torulose; style 0.5–1 mm; ovules and seeds 40–80 per fruit. Seeds brown, oblong to ovate, 0.7–
1.1 × 0.5–0.8 mm, wingless. 2n = 32, 64.
Flowering: Apr–Jul.
Habitat: marshes, steams, swamps, ditchts, seepage, springs, lake margins, mesic bottomland and
upland forests, wet areas, ledges of sheltered bluffs, on banks and in shallow water of streams
and spring branches, margins of crop fields, and waste ground.
Elevation: 0–2750 m.
Distribution: Canada (Alberta, British Columbia, Labrador, Manitoba, New Brunswick,
Newfoundland, Northwest Territories, Nova Scotia, Ontarioa, Prince Edward Island,
Quebeck, Saskatchewan, Yukon), United States (Alabama, Alaska, Arkansas, Clifornia,
Colorado, Connecticut, Delaware, District of Columbia, Florida, Georgia, Idaho, Illinois,
Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan,
Minnesota, Mississippi, Missouri, Montana, Nevada, New Hampshire, New Jersey, New
Mexico, New York, North Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania,
Rhode Island, South Carolina, Tennessee, Texas, Utah, Vermont, Virginia, Washington, West
Virginia, Wisconsin, Wyoming).
Cardamine pentaphyllos (Linnaeus) Crantz, Cl. Crucif. Emend. 127. 1769; Dentaria
pentaphyllos Linnaeus, Sp. Pl. 2: 654. 1753. TYPE: Dentaria heptaphyllos Bauhin in Horto
Dei Galloprovinciae, Herb Burser no. XVIII(1): 80 (lectotype designated by Hedge in Jarvis et
al. (1993: 42), UPS).
Cardamine penzesii Ančev & Marhold, Ann. Bot. Fenn. 36: 172. 1999. TYPE:
90
Cardamine picta W. J. Hooker, London J. Bot. 6: 292. 1847; Porphyrocodon pictum (W. J.
Hooker) J. D. Hooker in Bentham & J. D. Hooker, Gen. Pl. 1: 79. 1862. TYPE: Colombia,
New Granada, Paramo of Ruiz, Mar 1846, W. Purdie (holotype, K!; isotype, K!).
Herbs, perennial. Rhizomes slender, not scaly, with long internodes and nontuberous nodes.
Stems (3−)4.5−11 dm, erect or decumbent, simple or branched above, glabrous or rarely
pubescent. Cauline leaves (lower and middle) (7−)10−21(−25) cm, 7−11-foliolate; petiole
2.5−12 cm logn; lateral leaflet (1−)2−4(−5) × (0.7−)1.4−3 cm, broadly ovate to oblong-ovate,
glabrous or sparsely pubescent, base strongly oblique-subcordate or rarely cuneate, margin
incised-serrate, incised-crenate, or serrate, ciliolate with trichomes to 0.2 mm, teeth mucronate,
apex acute; petiolule 0.5−2.2 cm; terminal leaflet about same size as lateral; uppermost leaves
gradually smaller. Racemes bracteate throughout or rarely only basally; bracts compound,
gradually reduced in size and number of leaflets upward; flowers prominently protogynous;
fruiting pedicels (2−)2.5−4.2(−5) cm, divaricate-ascending, straight, stout. Sepals dark purple,
broadly oblong, 6−7 × 2.5−3.5 mm, glabrous; petals dark purple, broadly obovate, 1.5−2.2 ×
0.7−1 cm, base broadly cuneate, apex rounded to emarginated; filaments 6−7 mm; anthers
oblong, 1.5−1.7 mm; ovules 12−16 per ovary. Fruits 4.5−7 cm × 2−2.5 mm; style slender,
(6−)8−12(−20) mm. Seeds oblong, mm.
Flowering: Dec−Apr(−Aug).
Habitat: deep moss under Polylepis, paramo.
Elevation: 3300−3970 m.
Distribution: Colombia (Antioquia, Cauca), Ecuador (Pichincha).
Specimens examined: COLOMBIA. Antioquia: Termales, Páramo de Herveo, 3680 m,
1851−1857, Triana s.n. (G, P, W); Mpio de Urrao, between Páramo de Frontino and Finca La
Granada below El Quince, 6°25′N, 76°05′W, MacDougal, Roldán & Betancur 4585 (MO).
Cauca: Cordiller Central, Cabeceras de río Palo, quebrada del río López, Alto de Duende,
Cuatrecasas 19036 (F). ECUADOR. Pichincha: Páramo de Guamaní, Pifo-Papallacta rd,
78°13′W, 00°18′S, Øllgaard 98789 (AAU); Pifo-Papallacta rd, ca 1 km W of Paso de la Virgen,
78°13′W, 00°19′S, Laegaard 102330 (AAU); 1 km before Pasco de La Virgen, 78°11′W,
00°209′S, Laegaard 101371 (MO); Páramo de Guamani, Asplund 9643 (B, K, NY, S); Par.
Papallacta, Prescott 571 (DS, NY).
Seven sheets at K, all collected by Perdue from Paramo of Ruiz in New Granada, are authentic
material of Cardamine picta. However, as indicated in the original description, the sheet collected on
March 1846 should serve as the holotype.
Cardamine plumieri Villaris, Prosp. Hist. Pl. Dauphiné 38. 1779. TYPE: [France, Dauphiné],
“Grande Chartreuse” (holotype, GRM).
Cardamine pratensis Linnaeus, Sp. Pl. 2: 656. 1753; Dracamine pratensis (Linnaeus)
Nieuwland, Amer. Midl. Naturalist, 4: 40. 1915. TYPE: Herb. Linn. 835.15 (lectotype
designated by Khatri (1989: 92), LINN).
Herbs perennial, glabrous or rarely sparsely pilose basally. Rhizomes not fleshy, cylindrical,
not fragile; stolons absent. Stems (0.8–)1.5–5.5(–8) dm, erect, simple. Leaves thin, with raised
veins; rhizomal leaves to 30 cm; petiole (1–)2–7(–10) cm; terminal leaflet orbicular or broadly
obovate, 0.3–2 cm in diam., with a petiolule to 1.5 cm, base rounded, rarely subreniform or
cuneate, margin repand, apex rounded; lateral leaflets (0 or)2–8(–15) on each side of rachis,
91
about as large as or smaller than terminal leaflet, orbicular, ovate, or obovate, petiolulate or
sessile, margin crenate or repand; cauline leaves 2–12(–18), including petiole 2–17 cm,
pinnatisect, glabrous; petiole base not auriculate; terminal lobe linear, oblong, ovate, or
lanceolate, 1–2.5(–3.5) cm × 5–8(–10) mm, petiolulate or sessile; lateral lobes or leaflets 4–7(–
13) on each side of rachis, similar to terminal lobe, base petiolulate or sessile and decurrent,
margin entire or rarely dentate. Racemes ebracteate; fruiting pedicels erect-ascending or
subdivaricate, (0.5–)1.2–2.5(–3) cm. Sepals oblong or ovate, (2.5–)3–5(–6) × 1–2 mm, erect or
spreading, base saccate; petals purple or lilac, rarely white, obovate, (6–)8–15(–18) × 3–7.5(–10)
mm, clawed, apex rounded or emarginate; median filament pairs 5–10 mm, lateral pair 3–6 mm;
anthers narrowly oblong, (0.8–)1.2–2 mm. Fruit linear, (1.6–)2.5–4.5(–5) cm × (1.2–)1.5–2.3
mm; valves glabrous; style (0.5–)1–2.2(–2.7) mm, stout; ovules and seeds 20–30 per fruit. Seeds
light brown, oblong, 1.2–1.8(–2) × 1–1.4 mm. 2n = 16.
Flowering: May–Aug.
Habitat: Moist grounds, river or streamsides, limestone shores, sedge and grass meadows,
marshy pond margins, mossy areas, wet hollow, boggy areas, turfy shores, damp creek banks,
swamps, brooks and ditches, moist ravines, springy swales.
Elevation: 0–1000 m.
Distribution: all Europe, Canada (British Columbia, Newfoundland, New Brunswick, Nova
Scotia, Ontario, Quebec), China (Heilongjiang, Nei Mongol, Xinjiang, W Xizang), Japan,
Kazakhstan, Korea, Mongolia, Russia (European Part, Siberia, Far East), United States
(Connecticut, Indiana, Maine, Massachusetts, Michigan, New Hampshire, New Jersey, New
York, Ohio, Pennsylvania, Vermont).
Cardamine prorepens Fischer ex de Candolle, Syst. Nat. 2: 256. 1821; C. pratensis Linnaeus
var. prorepens (Fischer ex de Candolle) Maximowicz, TYPE: “Hab. in Siberia,
transbaikalensi, ad fl. Ingodem, c. Doroninsk in Dahuria, Fischer” (holotype, LE).
Cardamine borealis Andrzejowski ex de Candolle,
Cardamine pilosa Willdenow,
Cardamine pubescens Steven,
Herbs, perennial, glabrous or pubescnt. Rhizomes prostrate, stoloniferous. Stems 1.5–5(–6)
dm, stout, erect or decumbent, simple, glabrous or pubescent, 2–5(–8)-leaved. Rhizomal leaves
3–11-foliolate, 2–12 cm; petiole (0.5–)1.5–8(–10) cm; terminal leaflet ovate, oblong, or narrowly
obovate, 0.5–2(–3) cm × 4–13(–20) mm, with a petiolule 2–10 mm, margin crenate to repand;
lateral leaflets 1–5 pairs, often sessile, similar to terminal ones but smaller; cauline leaves 3–5foliolate, often glabrous; petiole to 7 cm, not auriculate at base; terminal leaflet elliptic, oblong,
or rhomboid, 1–3.5(–5) × 0.5–1.8(–2.5) cm, with a petiolule to 2.5 cm, base cuneate to obtuse,
margin remotely crenate, dentate, or repand, apex obtuse; lateral lobes obliquely ovate or oblong,
sessile or shortly petiolulate, similar to terminal one. Racemes ebracteate, 5–15-flowered;
fruiting pedicels 0.5–3 cm. Sepals ovate, 3–4.5 × 1–2 mm; petals white, narrowly obovate, 0.9–
1.5 cm × 5–7 mm, base cuneate, apex rounded or subemarginate; median filament pairs 5–7 mm,
lateral pairs 3–5.5 mm; anthers oblong, 1–1.8 mm; ovules 12–16 per ovary. Fruit linear, 1.5–4
cm × 1.5–2 mm; valves flat, glabrous or pilose; style slender, 2–4 mm. Seeds brown, ovate to
suboblong, 1.5–2.2 × 1–1.6 mm, wingless.
Flowering: May–Jul.
Habitat: river or stream banks, meadows.
Elevation: 1000–1700 m.
Distribution: China (Heilongjiang, Jilin, Nei Mongol), Korea, Mongolia, Russia (Far East).
92
Cardamine pulchella (J. D. Hooker & Thomson) Al-Shehbaz & G. Yang, Harvard Pap. Bot.
3(1): 77. 1998; Loxostemon pulchellus J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5:
147. 1861. TYPE: “Sikkim graminosis humidis! alt. 10,000−13,000 ped.,” J. D. Hooker, s.n.
(holotype, K!).
Loxostemon incanus R. C. Fang ex T. Y. Cheo & Y. Z. Lan, Bull. Bot. Res. North-East. Forest
Inst. 1(3): 52. 1981. TYPE: China, Yunnan, Chungtien Plateau, 7 Jul 1930, K. M. Feng 1559
(holotype, KUN!; listed as HKA).
Herbs, perennial, hirsute or glabrescent. Rhizomes to 1 cm, with stolons and numerous
bulbils; bulbils whitish, fleshy scales ovoid or subglobose, with rudimentary apical appendages.
Stems (5–)8–15(–20) cm, simple, erect; underground proximal part slender, glabrous;
aboveground part green or purplish, slender or stout, pilose or glabrous; bulbils of leaf axils
ovoid, to 3 × 2 mm. Rhizomal leaves 1(or 2), (1.5–)3–7 cm; petiole (1–)2.2–6 cm; terminal
leaflet broadly ovate or oblong, 3–10 × 2–4(–7) mm, with a petiolule to 3 mm; lateral leaflets
1(or 2) pairs, similar to terminal one. Cauline leaves 1–3, 1.2–5 cm; petiole 0.5–3.5 cm, base not
auriculate; terminal leaflet oblong or narrowly elliptic, (4–)6–12(–15) × (1–)1.5–4(–6) mm,
ciliate or glabrous, base cuneate, margin entire, apex mucronate; lateral leaflets 1(or 2) pairs,
similar to terminal one. Racemes ebracteate, 2–4(or 5)-flowered; fruiting pedicels ascending, 5–
13 mm, straight. Sepals ovate, 1.5–3 × 1–1.5 mm, margin membranous; petals deep purple or
mauve, broadly obovate, 5–8 × 2–4 mm, apex rounded. Median filament pairs 2–3 × 0.7–1.1
mm, flattened, extended apically into a tooth; lateral pair slender, 1–2 mm; anthers oblong, 0.5–
0.6 mm; ovules 12–16 per ovary. Fruit linear, 1–1.7 cm × 1–1.2 mm; valves smooth, glabrous;
style 0.5–1 mm. Seeds brown, broadly oblong, 1.3–1.6 × 0.8–1.1 mm, wingless.
Flowering: May–Aug.
Habitat: grassy marshlands, moist rocky places, stony streamsides, scree, mountain slopes.
Elevation: 3400–4600 m.
Distribution: Bhutan, China (Qinghai, Sichuan, Xizang, Yunnan), India (Sikkim), Nepal.
Cardamine purpurascens (O. E. Schulz) Al-Shehbaz, T. Y. Cheo, L. L. Lou & G. Yang, Novon
10: 324. 2000; C. microzyga O. E. Schulz var. purpurascens O. E. Schulz, Notizbl. Bot. Gart.
Berlin-Dahlem 11: 225. 1931; Loxostemon purpurascens (O. E. Schulz) R. C. Fang ex Y. C.
Lan & T. Y. Cheo, Bull. Bot. Res., Harbin 1(3): 54. 1981. LECTOTYPE (designated by AlShehbaz, 2000): China, SW Sichuan: Kulu Mountains, E of Muli Gomba, 4300 m, Jun 1928,
J. R. Rock 16487 (hololectotype, B!; isolectotypes, E!, MO!, US!).
Herbs, perennial, pilose. Rhizomes stout, 0.5–1.5 cm, often with several stolons. Stems
(8–)10–25(–30) cm, erect, simple, ridged, pilose, not flexuous. Basal leaves rosulate, 5–10 cm;
petiole 1–3.5 cm, ciliate; terminal leaflet subreniform or orbicular, 4–8 × 4–10 mm, petiolule 1–3
mm, base rounded or cordate, margin entire or obscurely and obtusely 3–5-lobed; lateral leaflets
3–7 pairs, obovate or suborbicular, symmetric or not, slightly smaller than terminal lobe, entire
or obscurely toothed, apex rounded; cauline leaves 2–10, 1–5 cm; petiole (2–)4–10(–15) mm,
ciliate, not auriculate at base; terminal leaflet linear, oblong, or lanceolate, 3–9 × 0.5–2 mm,
sessile or with a petiolule to 1.5 mm; lateral leaflets 4–7 pairs, narrowly oblong to oblong-ovate,
subequaling terminal leaflet, symmetric or not, pilose, base oblique or cuneate, proximal margin
entire or 1-toothed, distal margin entire, apex acute. Racemes ebracteate, many flowered; fruiting
pedicels divaricate or ascending, (0.7–)1–2.2(–3) cm, slender, pilose. Sepals oblong or ovate, 3–
4 × 1.5–2 mm, pilose, margin membranous, base of lateral pair saccate; petals magenta-red,
93
purple, or lavender, broadly obovate, 7–11 × 3–6 mm, cuneate into a clawlike base to 2 mm,
apex rounded; median filament pairs 4–5 mm, flattened below anther, to 1 mm wide; lateral pair
2.5–3.5 mm; anthers narrowly oblong, 0.9–1.2 mm; ovules 10–16 per ovary. Fruit 1.6–2.5 cm ×
1.2–1.8 mm; valves glabrous, smooth; style 1–2 mm. Seeds brown, broadly oblong, 1.2–1.5 ×
0.9–1 mm, wingless.
Flowering: May–Jul.
Habitat: river banks, wood margins, marshy places, swampy meadows, Rhododendron scrub.;
Elevation: 3500–4400 m.
Distribution: China (Sichuan, Yunnan).
Cardamine purpurea Chamisso & Schlechtendal, Linnaea 1: 20. 1826. TYPE:
Cardamine purpurea var. albiflos Hultén, Ark. Bot. (n.s.) 7: 62. 1968. TYPE:
Cardamine purpurea var. lactiflora O. E. Schulz ex Steffen, Beih. Bot. Centralbl. 54, abt. B:
551. 1936. TYPE:
Herbs, perennial, often cespitose, hirsute. Rhizomes not fleshy, vertical, 1–3 mm in diam.;
stolons absent. Stems (0.3–)0.5–1.2(–1.5) dm, most erect, 1 to several from base, simple, hirsute.
Basal leaves sometimes rosulate, 3–5-foliolate or rarely simple, (1.5–)2.5–7 cm, hirsute; petiole
(1.2–)2–5 cm; terminal leaflet reniform or suborbicular to broadly ovate, 3–10 × 4–15 mm, base
subcordate to rounded, margin entire to repand or obscurely 2-toothed, petiolule 0.5–6 mm;
lateral lobes distinctly smaller, sessile; cauline leaves 1–3, similar to basal leaves but smaller,
compound or rarely simple; petiole (0.2–)0.5–2, base not auriculate. Racemes ebracteate; fruiting
pedicels suberect ot ascending or divaricate, 5–12 mm, pubescent. Sepals oblong, 2–3(–4) × 1.4–
1.7 mm, erect, base of lateral pair not saccate; petals purple to pink or rarely white, obovate, 5–
7(–9) × 3–4(–5) mm, clawed, apex rounded; filaments of median stamens 2.5–3.5 mm, lateral
filaments 1.5–3.5 mm; anthers oblong, 0.6–0.9 mm. Fruit linear, 1.5–2.5 cm × 1.5–1.8 mm;
valves glabrous; style 1–2.5 mm; ovules and seeds 10–14 per fruit. Seeds brown, oblong to
broadly ovate, 1.7–2.1 × 1.4–1.7 mm, wingless. 2n = 96.
Flowering: Jun–Aug.
Habitat: moist tundra, damp woods and ravines, alpine turf, river flats, peaty subarctic meadows,
streamsides, moist slopes.
Elevation: 600–1800 m.
Distribution: Canada (Yukon), Russia (Far East), United States (Alaska).
Cardamine quinquefolia (M. Bieberstein) Schmalhausen, Fl. Sredn. Juž. Rossii 1: 51. 1895;
Dentaria quinquefolia M. Bieberstein, Fl. Taur.-Caucas. 2: 109. 1808. TYPE : Ex Tauria et
Caucaso, [F. A. Marshall von Bieberstein] (lectotype designated by Marhold (2001: 45), LE,
plant on upper right corner of sheet).
Cardamine raphanifolia
Cardamine repens (Franchet) Diels, Notes Roy. Bot. Gard. Edinburgh 5: 204. 1912; Dentaria
repens Franchet, Bull. Soc. Bot. France 32: 5. 1885; Cardamine tenuifolia (Ledebour)
Turczaninow var. repens (Franchet) Franchet, Bull. Soc. Bot. France 33: 399. 1886;
Loxostemon repens (Franchet) Handel-Mazzetti. TYPE: China, Yunnan: San-tchang-kiou,
secus viam a Tali and Hokin, 27 May 1884, Delavay 65 (holotype, P!).
Herbs, perennial, glabrous throughout except for apices of leaf lobes. Rhizomes much
elongated, with a few bulbils, 3–8 mm wide. Stems 10–45 cm, simple, erect, straight, stout.
Rhizomal leaves simple or trifoliolate; petiole 4–8 cm; leaf blade or terminal leaflet suborbicular
94
or broadly cordate, to 1.5 cm in diam., base cordate, margin entire or obscurely 5-lobed, terminal
leaflet with a petiolule to 7 mm; cauline leaves 2–5, trifid or pinnatisect and with 2(or 3) lateral
lobes on each side of midvein, lobes all decurrent, often 1- or 2-toothed, rarely entire; petiole
0.3–4 cm, winged, glabrous or ciliate, not auriculate or rarely with a small, toothlike, puberulent
auricle to 0.3 mm; terminal lobe linear-lanceolate, 1.2–5 cm × 4–9 mm, base attenuate, margin
entire, along distal half often minutely puberulent with stout trichomes rarely to 0.2 mm, apex
acute; lateral lobes similar to terminal. Racemes ebracteate, 5–20-flowered; fruiting pedicels
ascending, 0.8–2.2 cm, straight, glabrous. Sepals ovate, 2.5–3 × 1.2–1.7 mm, glabrous, margin
and apex membranous, lateral pair subsaccate; petals white or lavender, broadly obovate or
spatulate, 6–8 × 2.5–4 mm, apex rounded; median filament pairs 2.5–4, slightly flattened; lateral
pair 1.5–3 mm; anthers oblong, 0.8–1 mm; ovules 8–14 per ovary. Fruit linear, 2–3 cm × ca. 1.5
mm; valves smooth, glabrous; style 1–5 mm. Seeds brown, oblong, 1.2–1.5 × ca. 0.7 mm,
wingless.
Flowering: May–Jun.
Habitat: grassy slopes, moist rocky crevices.
Elevation: 2400–3400 m.
Distribution: China (Sichuan, Yunnan).
Cardamine resedifolia Linnaeus, Sp. Pl. 2: 656. 1753. TYPE: illustration in Bauhin (Prodromus
theatri botanici 1620: 45) captioned “Nasturtium alpinum minus Resedae foliis” (lectotype
designated by Marhold (1996: 125). Epitype designated by Marhold (1996: 125); Pyrenaen
orient.: Porté Puymorens - Vallée de Lanous, E und W Col. de Lanoux, c. 2400 m, im
Silikatschutt, 4-17 Aug 1974, A. Polatschek s.n. (W).
Cardamine rivularis
Cardamine robusta I. Thompson, Muelleria 9: 151. 1996. TYPE: Australia, New South Wales,
Club Lake, Kosciusko area, 36º25′S, 148º16′E, 10 Jan 1960, B. G. Briggs s.n. (holotype,
NSW).
Herbs, pererrnial, glabrous, forming dense clusters to 1 m wide. Rhizomes slender, frequently
branched. Stems to 30 cm, slender, branched below, glabrous. Basal leaves rosulate, somewhat
fleshy, long petiolate, to ca. 25 cm, 3–5(–7)-foliolate; terminal leaflet broadly ovate to oblong,
base cuneate to slightly cordate; lateral leaflets smaller than and same shape as terminal leaflet,
long petiolulate; cauline leaves 0–4, petiolate, not auriculate at base, similar to but smaller than
basal leaves, with narrower leaflets, or divided and not compound. Racemes ebracteate, few to
many flowered, condensed or sometimes elongated; fruiting pedicels stout, to 20 mm. Sepals
ovate, 3–4.5 mm; petals white, differentiated into blade and claw, 8–12 × 5–7 mm; stamens 6.
Fruits linear, 2–4 cm × 2–3 mm; style 1–3 mm. Seeds oblong-elliptic, 2–2,5 mm.
Flowering: Jan–Apr.
Habitat: alpine regions among granite boulders, moist slopes bordering melting snow, shores of
glacial lakes.
Distribution: Australia (New South Wales).
Cardamine rockii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 9: 473. 1926. TYPE: China,
SW Sichuan, Muli or Muli Kingdon, 10,000–14,000 m, Jun 1922, J. F. Rock 5585 (holotype,
B!; isotypes, E!, GH!, P!, US!, W!).
Herbs, perennial, pilose or subhirsute. Rhizomes elongated, 1–3 mm in diam. Stems (9–)15–
45(–55) cm, decumbent basally, erect, simple, ridged, pilose, 4–10-leaved. Basal leaves not
95
rosulate, similar to cauline ones; lower and middle cauline leaves (1.5–)3–12(–15) cm; petiole
0.5–4 cm, ciliate, not auriculate at base; terminal leaflet broadly ovate or suborbicular, (3–)5–
12(–15) × (2–)4–8(–10) mm, symmetrical, with a petiolule 1–3 mm, base obtuse, truncate, or
cuneate, margin 2–4-toothed on each side, teeth submucronate; lateral leaflets alternate or rarely
opposite, 6–12 on each side of rachis, broadly to narrowly ovate or ovate-lanceolate, subequaling
terminal leaflet, asymmetric, sessile or with a petiolule to 3 mm, pilose or rarely subglabrous,
base oblique, proximal margin coarsely and unequally (2 or)3–5(or 6)-toothed, distal margin
entire or 1–3-toothed, acute apex and teeth submucronate. Racemes ebracteate, many flowered;
flowering pedicels divaricate or ascending, 1–2 cm, slender, pilose. Sepals oblong to narrowly
ovate, (2.5–)3–4 × (1–)1.2–1.6 mm, margin membranous, base of lateral pair subsaccate; petals
white, obovate, 7–9 × 2.5–4 mm, cuneate into a clawlike base to 2 mm, apex rounded or
subemarginate; median filament pairs 4–5 mm, slightly dilated, not flattened; lateral pair 2.5–3.5
mm; anthers narrowly oblong, 1–1.5 mm; ovules 10–14 per ovary. Fruit and seeds not seen.
Flowering May–Jul.
Habitat: moist grassy areas, boggy places by streams, alpine meadows.
Elevation: 3100–4700 m.
Distribution: China (Sichuan, Yunnan).
Cardamine rostrata Grisebach, Abh. Königl. Ges. Wiss. Göttingen 6: 115. 1854. TYPE: Chile,
Cordillera de Ranco. Lechler 841 (holotype, GOET, not seen, fragments, BAA!; isotypes, G!,
K!, 3P!, W!).
Cardamine reniformis Philippi, Anal. Univ. Chile 27(2): 313. 1856; C. rostrata var. reniformis
(Philippi) O. E. Schulz, Bot. Jahrb. Syst. 32: 434. 1903. TYPE: Chile, Valdivia: en el lugar
llamado Hualleria, Oct 1864, F. Philippi 55 (holotype, SGO-63900!; fragments, BAA!). The
type information was copied from Muñoz (1960) who listed also SGO-49343 as part of the
type collection. I have not seen the latter sheet.
Cardamine lechleriana Steudel, Flora 39: 409. 1856. TYPE: Chile, Corillera de Ranco, Lechler
2249 (holotype, P!; isotypes, ).
Cardamine rostrata var. dichondroides Spegazzini, Anal. Mus. Nac. Buenos Aires 7: 212. 1902.
TYPE: Argentina, Dep. Languiñeo, Río Chubut, Dec 1900, N. Illin s.n. (holotype, LP #…..).
Herbs, perennial, glabrous throughout. Rhizomes not tuberous, rather slender, not scaly.
Stems often 1 from rhizome, (8−)15−45(−60) cm, erect but subdecumbent at base, glabrous,
simple or rarely branched above, remotely 3−7-leaved. Basal leaves not rosulate, they and
lowermost cauline leaves simple or rarely with 1 or 2 pairs of lateral leaflets; petiole 2−7(−11)
cm; leaf blade or terminal leaflet orbicular to cordate or reniform, (1−)1.5−3.5(−4) ×1.8−4(−5)
cm, often wider than long, palmately veined, not fleshy, base often deeply cordate, 5−9-angled,
the vein-ending angle terminating in a mucro, rarely margin repand; lateral leaflets (when
present) on petiolules 1−3 mm, similar in shape and margin and considerably smaller than
termial leaflet, base usually oblique; middle and upper cauline leaves simple or rarely, 3−5foliolate leaflets, smaller and narrower than lowermost leaves. Racemes ebracteate, corymbose,
elongated considerably in fruit; rachis straight; fruiting pedicels (0.5−)0.8−1.7(−2) cm, ascending
to divaricate, straight and often forming a straight line with fruit, glabrous. Sepals oblong, 2.5−4
× 1−1.5 mm, erect, caducous, glabrous; petals white, oblanceolate-obovate, (6−)7−10(−12) ×
2.5−5 mm, apex obtuse; filaments 3.5−5 mm; anthers oblong, 0.7−1 mm; ovules 12−20 per
96
ovary. Fruits linear, (1.8−)2.5−4 cm × 1.2−1.7 mm, attenuate to apex; style slender
(1.5−)2−5(−6) mm. Seeds brown, oblong to broadly ovate, 1.4−1.7 × 0.9−1.2 mm.
Flowering: Oct−Mar.
Habitat: Nothofagus woods, moist areas.
Elevation: 0−1600 m.
Distribution: Chile (Región VIII, IX, and X).
Specimens examined: CHILE. Cautin, Faldeos de Volcan Llaima, Muñoz 6415 (SGO), Volcan
Llaima, Montaldo & Seeger 165 (SGO); Malleco, Dep. Curacautin, Cerros al N de
Malalcahuello, Montaldo 386 (SGO). VIII: Dep. San Carlos, Camino de San Fabián de Alico a
los Sauces, 4 km ade Los Sauces, [36°38′S, 71°24′W], Weldt 486 (CONC); Cordillera de
Nahuelbuta, Bajo Pino Huacho, [37°43′S, 73°06′W], Marticorena, Quezada & Rodríguez 1719
(CONC); Reerva Forestal Pino Huacho, [37°42′S, 73°09′W], Marticorena, Quezada & Rodríguez
1600 (CONC); Parq Nac. Laguna del Laja, 37°25′S, 71°25′W, Godoy 341 (CONC). IX: Volcán
Llaima, [38°43′S, 71°43′W], 30 Jan. 1942, Gunckel s.n. (CONC), Montero 4786, 4895 (CONC),
Sparre 4887 (SGO), Garaventa 4520 (BACP, SI), Garaventa 4521 (BACP, CONC, SI); Dep. t.
Victoria, Fundo San Elías, Sparre 3292 (SGO); rd to Laguna Malleco, Salto de la Culebra and
Laguna Verde, Morrison & Wagenknecht 17531 (DS, UC); Camino de Curacautin a Lonquimay,
km 46, [38°27′S, 71°26′W], Ricardi & Marticorena (CONC); Malalcahuello, [38°28′S,
71°35′W], Zöllner 7602 (CONC), Zöllner 8781 (MO); Camino de Termas de Manzanares a
Lonquimay, km 26, [38°25′S, 71°25′W], Ricardi & Marticorena 5009/1393 (CONC); Orillas del
Río Niblinto, [38°10′S, 71°45′W], Marticorena & Quezada 1531 (CONC); Paso de Lonquimay,
[38°22′S, 71°34′W], Valle de Lonquimay, Montaldo 4546 (CONC); Zöllner 8142 (CONC, NA);
Termas de Tolhuaca, [38°14′S, 71°44′W], Gunckel 16367 (CONC, US), Looser 2660 (BACP,
SI); Curacautin, Cord. de Las Raices, [38°26′S, 71°29′W], Montero 10325 (CONC); Moleau
Mocho, [39°56′S, 72°04′W], Montero 4994 (CONC); Cautín, Laguna Verde, Parq. Nac.
Huerquehue, 39°08′S, 71°43′W, Godoy 414 (CONC). X: Dep. Unión, La Guallería, Sparre 3691
(S, SGO); Gualleria (as Hualleria), Philippi s.n. (B, SGO, W); Cumleufo (Trumao), [40°21′S,
73°10′W], Hollermayer 637 (CONC, SI); Lago Rinihue, [39°54′S, 72°10′W], Montero 5036
(CONC); Arededores de Osorno, [40°35′S, 73°08′W], Seki 82 (CONC); Chiloe, Piruquina,
[42°24′S, 73°48′W], Junge 79 (CONC).
In the overall aspects of the plant, Cardamine rostrata resembles C. tuberosa, but it can be
easily separated from that by the non-tuberous, slender rhizomes and often angled leaves. It also
resembles C. codata, but differs in having non-fleshy leaves, angled instead or crenate leaves, and
lax inflorescence consipicuously expanded in fruit.
Cardamine rotundifolia Michaux, Fl. Bor.-Amer. 2: 30. 1803; Dentaria rotundifolia (Michaux)
Greene, Pittonia 3: 124. 1896. TYPE:
Herbs, perennial, glabrous throughout. Rhizomes slender, to 2 mm in diam; stolons absent.
Stems (1–)1.5–3(–5) dm, most commonly procumbent, sometimes erect, simple or branched, not
flexuous, glabrous. Rhizomal leaves absent, basal leaves not rosulate, soon withered; cauline
leaves simple; petiole (0.3–)0.5–2.5(–4) cm; leaf blade of lower and middle ones oblong, ovate,
suborbicular, or cordate, (0.5–)1–4.5(–5.5) × (0.5–)1–4(–5.4) cm, glabrous, base cordate,
rounded, or truncate, margin, entire, repand, or sinuate; upper leaves gradually reduced in size
and with shorter petioles. Racemes ebracteate; fruiting pedicels divaricate to ascending, (0.6–)1–
97
1.5(–2) cm. Sepals oblong, 2–3 × 1–1.7 mm, erect, base of lateral pair not saccate; petals white,
broadly oblanceolate, spreading, 5–7(–8) × 2–3 mm, not clawed, apex rounded; filaments of
median stamens 3.5–4.5 mm, lateral filaments 2.7–3.5 mm; anthers oblong, 0.8–1.2 mm. Fruit
linear, 1–1.5(–2) cm × 0.8–1.1 mm; valves glabrous, torulose; style 1.5–2.5 mm; ovules and
seeds 40–80 per fruit. Seeds brown, oblong to ovate, 0.9–1.1 × 0.5–0.6 mm, wingless.
Flowering: Apr–Jun.
Habitat: streambanks, swamps, low woodlands, wet rocky areas, seepage areas.
Elevation: 150–400.
Distribution: United States (Delaware, Georgia, Kentucky, Maryland, New Jersey, New York,
North Carolina, Ohio, Pennsylvania, Tennessee, Virginia, West Virginia).
Cardamine rupicola (O. E. Schulz) C. L. Hitchcock, Univ. Wash. Publ. Biol. 17(2): 474. 1964;
C. californica Nuttall var. rupicola O. E. Schulz, Bot. Jahrb. Syst. 32: 388. 1903; Dentaria
rupicola (O. E. Schulz) Rydberg, Fl. Rocky Mts. 348. 1917. TYPE:
Herbs, perennial, glabrous throughout. Rhizomes slender, cylindrical, 1–2 mm in diam.;
stolons absent. Stems 0.6–2 dm, erect or decumbent at base, simple, glabrous. Rhizomal leaves
palmately or subpalmately compound, 3–5(–7)-foliolate, 5.5–17(–22) cm, fleshy; petiole 4–14(–
17) cm; terminal leaflet ovate to lanceolate or elliptic-oblong, 1–3 × 0.6–1.7 cm, with a petiolule
1–5 mm, glabrous, base cuneate or obtuse, margin entire, apex apiculate; lateral leaflets about as
large as or smaller than terminal leaflet, subsessile, margin same as terminal leaflet; cauline
leaves 2 or 3, 3–5-foliotate; petiole 0.7–4(–8) cm, base not auriculate; terminal lobe elliptic to
oblong, or ovate, 1.2–3.5 × 0.4–2.5 cm, entire, petiolule 1–5 mm; lateral leaflets similar to
terminal one but smaller, sessile. Racemes ebracteate; fruiting pedicels ascending to divaricate,
0.6–1.8 cm. Sepals oblong, 3–5 × 1.5–2 mm, erect, base of lateral pair saccate; petals white,
obovate, 8–14 × 4–7 mm, short clawed, apex rounded or subemarginate; median filament pairs
4–5 mm, lateral pair 2.5–3.5 mm; anthers oblong, 1–1.2 mm. Fruit linear, 2–4 cm × 1.5–2.2 mm;
valves glabrous; style 1–5 mm; ovules and seeds 10–14 per fruit. Seeds brown, oblong, 1.8–2.2 ×
1.2–1.5 mm.
Flowering: Jul–Aug.
Habitat: limestone talus slopes, loose limy shale, moist banks.
Elevation: 2200–2700.
Distribution: United States (W Montana).
Cardamine scaposa Franchet, Pl. David. 1: 33. 1883. TYPE: China, Nei Mongol, Géhol, MayJun 1865, A. J. P. David 1815 (holotype, P!; isotypes, LE!, P!).
Cardamine denudata O. E. Schulz, Bot. Jahrb. Syst. 36(Beibl. 82): 46. 1905. TYPE: China,
Shaanxi. Sciu-ian-shan, Kanyhuo, 15 May 1899, G. Giraldi 5480 (holotype, B!).
Herbs, perennial, scapose, glabrous throughout. Rhizomes slender, with slender stolons.
Stems (4–)8–18 cm, leafless, erect, simple. Rhizomal leaves simple; petiole (1–)2–9(–12) cm;
leaf blade reniform or suborbicular, (0.3–)0.6–1.7(–2) × (0.5–)1–2.5(–3) cm, base cordate,
margin repand-crenate or entire; cauline leaves absent. Racemes ebracteate, 2–7(–10)-flowered;
fruiting pedicels erect or erect-ascending, 1–4 cm, proximal longest. Sepals ovate or oblong, 3–4
× 1.5–2.2 mm, margin membranous, lateral pair subsaccate; petals white, broadly obovate,
(0.8–)0.9–1.3 cm × 5–7 mm, cuneate into a clawlike base to 2 mm, apex rounded or
subemarginate; median filament pairs 4.5–8 mm, slightly dilated at base; lateral pair 2.5–4.5 mm;
anthers narrowly oblong, 1.5–1.8 mm; ovules 8–14 per ovary. Fruit linear, 2–3.5 cm × 1.2–1.7
mm; valves glabrous, smooth; style 3–7.5 mm. Seeds brown, oblong, 2–3 × 1–1.5 mm, wingless.
98
Flowering: Apr–Jun.
Habitat: shrubby slopes, moist areas.
Elevation: 1400–2500 m.
Distribution: China (Hubei, Shaanxi, Shanxi, Sichuan).
Cardamine schinziana O. E. Schulz, Bot. Jahrb. Syst. 32: 503. 1903; C. yezoensis Maximowicz
var. schinziana (O. E. Schulz) Ohwi, Fl. Jap. ed. 2, 1437. 1965. TYPE :
Cardamine nasturtiifolia H. Boissieu, Bull. Herb. Boiss. 7: 793. 1899; non Bertol. ex Steud.,
Nomencl. Bot. Ed. 2, 1: 281. 1840. TYPE: Japan, Forêt de Sarura, 1 Jul 1893, Père Urbain
Faurie 10482 (holotype, P; isotype, MO!).
Cardamine nasturtiifolia H. Boissieu var. lasiocarpa H. Hara, Bot. Mag. (Tokyo) 49: 72. 1935.
Cardamine schinziana O. E. Schulz var. lasiocarpa (H. Hara) Koidzumi, Acta Phytotax. Geobot.
5: 121. 1936.
Herbs, perennial, glabrous. Rhizomes creeping, short, somewhat thickened. Stems 20−50
cm, erect to subdecumbent, branched above. Lower leaves and middle cauline 8−15, 4−15 cm;
terminal leaflet obovate to orbicular, 1.5−2.5 cm, 1.3−2.1 cm wide, base truncate or obtuse,
margin crenate, apex rounded to obtuse; lateral leaflets 2−5 on each side of rachis, ellipticlanceolate to oblong, 0.4−1.8 cm, 0.3−1.3 cm wide, crenate or rarely entire; uppermost leaves
with 2−6 lateral leaflets on each side of rachis; terminal leaflet narrowly lanceolate, oblanceolate,
or suboblong, 0.8−3(−5) cm, 0.3−1.5 cm wide, base cuneate, margin entire or few toothed, apex
acuminate to acute; lateral leaflets lanceolate to elliptic, 0.8−3 cm, 5−10(−15) mm wide, coarsely
toothed or 1−4-shallowly lobed on each side. Raceme ebracteate; fruiting racemes 7−15 cm.
Fruiting pedicels (1−)1.2−1.7(−2) cm, strongly patent. Sepals 2.5−3.5 mm, 1−1.5 mm wide;
petals white, obovate, 6−9 mm, 3−4.5 mm wide; filaments of median staminal pairs 5−6 mm,
those of lateral pair 3.5−4.5 mm; anthers 1−1.5 mm; ovules 16−20 per ovary. Fruits linear,
(1.5−)2−3.5(−4) cm, 1.2−1.8 mm wide, glabrous or sparsely pubsecent; style slender,
(1−)1.5−3(−4) mm. Seeds light brown, oblong-ovate, 1.5−2.2 mm, 1−1.5 mm wide.
Flowering: Jun−Jul.
Habitat: wet places along streams, river banks.
Distribution: Japan (Hokkaido/Kurils), Russia (Far East).
Cardamine schulzii
Cardamine scutata Thunberg, Trans. Linn. Soc. London 2: 339. 1794; C. flexuosa Withering
var. scutata (Thunberg) O. E. Schulz, Bot. Jahrb. Syst. 32: 477. 1903. TYPE:
Cardamine angulata W. J. Hooker var. kamtschatica Regel, Bull. Soc. Imp. Naturalistes Moscou
34: 172. 1861; C. flexuosa Withering var. kamtschatica (Regel) Matsumura, publication
TYPE:
Cardamine autumnalis Koidzumi, Bot. Mag. (Tokyo) 43: 404. 1929. TYPE:
Cardamine baishanensis P. Y. Fu, Fl. Pl. Herb. China Bor.-Or. 4: 229. 1980. TYPE:
Cardamine dentipetala Matsumura, Bot. Mag. (Tokyo) 13: 51. 1899. TYPE:
Cardamine dentipetala var. longifructus (Ohwi) Hiyama, J. Jap. Bot. 38: 58. 1963. TYPE:
Cardamine drakeana H. Boissieu, Bull. Herb. Boiss. 7: 791. 1899. TYPE:
Cardamine hirsuta Linnaeus var. latifolia Maximowicz, Bull. Acad. Imp. Sci. Saint Pétersbourg
18: 279. 1873. C. flexuosa var. latifolia (Maximowicz) Makino, Shokubutsu-Dzukan 447.
99
1925; C. scutata var. latifolia (Maximowicz) H. Hara, J. Fac. Scil. Univ. Tokyo III, 6: 59.
1952. TYPE:
Cardamine regeliana Miquel, Ann. Mus. Bot. Lugduno-Batavi 2: 73. 1866. C. flexuosa
Withering subsp. regeliana (Miquel) O. E. Schulz, Bot. Jahrb. Syst. 32: 476. 1903; C.
flexuosa var. regeliana (Miquel) Matsumura, Bot. Mag. (Tokyo) 13: 73. 1899. C. hirsuta var.
regeliana (Miquel) Maximowicz, Bull. Acad. Imp. Sci. Saint Pétersbourg 18: 279. 1873; C.
scutata subsp. regeliana (Miquel) H. Hara, J. Fac. Sci. Univ. Tokyo, III, 6: 59. 1952; C.
sylvatica Link var. regeliana (Miquel) Franchet & Savatier, Enum. Pl. Jap. 1: 35. 1875.
TYPE:
Cardamine flexuosa Withering var. manshurica Komarov, Trudy Imp. St.-Peterburgsk. Bot. Sada
22: 269. 1903; C. regeliana Miquel var. manshurica (Komarov) Kitagawa, Lineam. Fl.
Mansh. 228. 1939. TYPE:
Cardamine hirsuta Linnaeus var. rotundiloba Hayata, J. Coll. Sci. Univ. Tokyo 30: 31. 1911; C.
scutata var. rotundiloba (Hayata) Liu & S. S. Ying, Fl. Taiwan 2: 686. 1976. TYPE:
Cardamine longifructus Ohwi, J. Jap. Bot. 33: 211. 1958. TYPE:
Cardamine scutata var. longiloba P. Y. Fu, Fl. Pl. Herb. China Bor.-Or. 4: 229. 1980. TYPE:
Cardamine taquetii H. Léveillé, Repert. Sp. Nov. Regni Veg. 8: 259. 1910. TYPE: Korea,
Quelpaert, Hongno, Oct 1908, Taquet 563 (holotype, E!; isotype, K!).
Cardamine zhejiangensis T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. North-East. Forest. Inst.,
Harbin 1980(6): 24. 1980. TYPE: China, Zhejiang, Xitianmu Shan, 4 May 1957, Ho Hseyu
20927 (holotype, HHBG; isotype, NAS!).
Cardamine zhejiangensis var. huangshanensis D. C. Zhang, Bull. Bot. Res., Harbin (11)1: 41.
1991. TYPE: China, Anhui: Huangshan, Tanglingguan, 900 m, 14 May 1989, Shao Jianzhang
89008 (holotype, ANU).
Herbs, annual or biennial, rarely short-lived perennial, glabrous or sparsely pilose. Stems
(5–)15–50(–70) cm, erect, simple at base, simple or branched above, not flexuous. Basal leaves
not rosulate, often withered by anthesis; petiole to 3.5 cm; leaf blade pinnatisect, with 1–4 lateral
lobes on each side of midvein; terminal lobe subreniform, suborbicular, rhomboid-ovate, or
broadly obovate, much larger than lateral ones, (1–)1.5–2.5 × 0.7–2 cm, repand, crenate, or 3–5lobed; lateral lobes petiolulate or subsessile, oblong, ovate, or suborbicular. Cauline leaves
similar to basal ones; petiole to 3 cm, base not auriculate; lateral lobes 1–5 on each side of
midvein, much smaller than terminal lobe; terminal lobe (0.7–)2–5(–6.5) × (0.5–)1.5–4(–5) cm,
repand, crenate, or coarsely 3–5(–7)-lobed. Racemes ebracteate; rachis straight; fruiting pedicels
divaricate or ascending, (0.3–)0.6–1.4(–1.8) cm, slender. Sepals oblong, 1.5–2.5(–3) ×
(0.7–)0.9–1.4 mm; petals white, spatulate, 2.5–4.5(–6) × (1–)1.5–2.5 mm; stamens 6; filaments
2–3.5 mm; anthers ovate, 0.3–0.6 mm; ovules 20–40 per ovary. Fruit linear, (0.9–)1.5–2.8(–3.5)
cm × (0.8–)1–1.4 mm; valves glabrous or rarely sparsely puberulent or pilose, torulose; style
(0.3–)0.6–1.5 mm. Seeds brown, oblong or subquadrate, 0.9–1.5 × 0.6–0.9 mm, narrowly
margined or not. 2n = 32.
Flowering: Apr–Jul.
Habitat: valleys, shady slopes, damp sites, along ditches, rock crevices, mountain slopes,
roadsides, streamsides.
Elevation: 0–2100 m.
Distribution: China (Anhui, Guangdong, Guizhou, Jiangsu, Jilin, Sichuan, Taiwan, Zhejiang),
Japan, Korea, Russia (Far East).
Cardamine seidltiziana
100
Cardamine seravschanica
Cardamine silana Marhold & Perný, Willdenowia 33: 69. 2003. TYPE:
Cardamine simplex Handel-Mazzetti, Symb. Sin. 7: 361. 1931. TYPE: China, Yunnan, above
Ganhaidse, Lidjiang, 3200 m, 22 Jul 1914, Handel-Mazzetti 4310 (holotype, WU!; isotypes,
E!, GH!, NAS!, W!).
Cardamine truncatolobata W. T. Wang, Acta Bot. Yunnan. 9: 14. 1987. TYPE: China, Sichuan,
Luding, Mons Gonggashan, Yanzigou, 3500–3800 m, 30 Jun 1982, Lan Kaiyong, Li
Liangqian & Fei Yong 341 (holotype, PE!; isotypes, KUN!, PE!).
Loxostemon axillus Y. C. Lan & T. Y. Cheo, Bull. Bot. Res., Harbin 1(3): 52. 1981. TYPE:
China, Sichuan, Tien-chuan-hsien, Chu 2528 (holotype, HNWP; isotypes, E!, NAS!, W!).
Herbs perennial, glabrous throughout or sparsely pilose. Rhizomes slender, thickened at stem
base, with 1 or few stolons. Stems (8–)13–35 cm, erect, slender, simple or 1- or 2-branched,
flexuous. Basal leaves 1–3, 3- or 5(or 7)-foliolate; petiole (1.3–)2–6(–8) cm; terminal leaflet
broadly obovate, 3–12 × 5–13 mm, with a petiolule 1–5(–8) mm, base subtruncate or rounded,
margin entire or apically subtruncate or obtusely 3-lobed, ultimately mucronate; lateral leaflets 1
or 2(or 3) pairs, subsessile or petiolulate, somewhat similar to terminal leaflet but smaller.
Cauline leaves (1 or)2–5(–7), 3- or 5-foliolate; petiole (0.8–)1.2–3 cm, not auriculate at base;
terminal leaflet of uppermost leaves filiform, linear, or rarely narrowly oblanceolate, 0.5–2 cm ×
(0.3–)1–2 mm, attenuate at base, margin entire, apex acute. Racemes ebracteate, lax, (2–)5–14(–
16)-flowered, rachis often flexuous; fruiting pedicels erect or ascending, (0.8–)1–2.5(–3) cm,
straight, glabrous, proximal ones often much longer than distal. Sepals ovate or oblong, 2.5–3.5
× 1.3–1.8 mm, glabrous, lateral pair subsaccate; petals white, obovate, (6.5–)7–9 × 3.5–5 mm,
not clawed, apex rounded; median filament pairs 4–5 mm, slender; lateral pair 3–3.5 mm; anthers
oblong, 0.8–1.1 mm; ovules 8–14 per ovary. Fruit linear, (1–)1.5–2.5(–2.8) cm × 1–1.2 mm;
valves glabrous, smooth; style (1–)1.5–3(–4) mm. Seeds brown, oblong, 1.2–1.6 × 0.8–1.2 mm,
wingless.
Flowering: May–Jul.
Habitat: meadows, damp turf, marshy areas, stream or ditch sides.
Elevation: 2500–3800 m.
Distribution: China (Sichuan, Yunnan).
Cardamine speciosa Britton, Bull. Torrey Bot. Club 16: 16. 1889; C. jamesonii Hooker var.
speciosa (Britton) O. E. Schulz, Bot. Jahrb. Syst. 32: 422. 1903. TYPE: Bolivia, La Paz,
Undavi, 1885, 10,000 ft, H. H. Rusby 1199 (holotype, NY!; isotype, NY!).
Herbs, perennial, with fleshy, elongated, non-scaly rhizomes. Stems 5−45 cm, erect or
decumbent only at base, simple or branched above, often glabrous. Rhizomal leaves and
lowermost cauline compound, pinnate, 2−15 cm; petiole 1−8 cm; lateral leaflets 2 or 3 pairs,
sessile or nearly so, broadly to narrowly obovate, 0.6−25 × 0.4−1.7 cm, sparsely to pubescent or
ciliate with trichomes 0.2−0.5 mm, base oblique or rarely cuneate, margin entire or obscurely 1toothed, apex obtuse, distinctly mucronate; terminal leaflet same size or slightly larger than
lateral ones, short petiolulate, obovate, entire or few toothed; cauline leaves 2−4, much smaller
than those of basal leaves; leaflets oblong to oblong-linear, much different in size and shape than
basal ones, entire, mucronate. Racemes ebracteate, many flowered, dense corymbose, slightly
101
elongated in fruit; rachis straight; fruiting pedicels 1−2.5 cm, stout, ascending, straight. Sepals
purple, oblong, 3−4.5 × 2−2.5 mm, caducous, glabrous; petals purple, broadly obovate, 0.9−1.2
cm × 4−6 mm, apex obtuse or rounded; filaments 5−7 mm; anthers oblong, 1.2−1.5 mm; ovules
…−… per ovary. Fruits linear, 4−5 cm ×..−… mm, glabrous; style ….−5 mm. Seeds brown,
oblong,..−… × …−… mm.
Flowering:.
Habitat: Wet mossy areas.
Elevation: 3000−3300 m.
Distribution: Bolivia (La Paz/Undavi).
Specimens examined: BOLIVIA. La Paz: Undavi, North Yungas, 3300 m, Nov. 1910, Buchtein
126 (E, F, G, GH, NY); Buchtien 585 (NY).
Schulz (1903) reduced Cardamine speciosa to a variety of C. jamesonii, but the two are quite
differenent morphologically and disjunct geographically. The latter is distributed in Ecuador,
Venezuela and Colombia, whereas C. speciosa is endemic to Bolivia, and the two taxa are
separated by more than 2000 air kilometers. Cardamine speciosa can easily be separated from C.
jamesonii by having ebracteate (vs. bracteate) racemes, entire or repand (vs. crenate, incisedcrenate, or serrate leaflets), 2−4 (vs. 5−10) cauline leaves excluding bracts, petiolulate (vs.
sessile) lateral leaflets, and leaflets of cauline leaves drastically narrower and smaller (vs. about
the same shape and size) as the lowermost leaves.
Cardamine sphenophylla
Cardamine stellata J. D. Hooker, Fl. Antarct.
Cardamine stenoloba Hemsley, J. Linn. Soc., Bot. 29: 303. 1892; Loxostemon stenolobus
(Hemsley) Y. C. Lan & T. Y. Cheo, Bull. Bot. Res., Harbin 1(3): 56. 1981. TYPE China, W
Sichuan, near Tachienlu [Kangding], A. E. Pratt 352 (lectotype designated by Cheo et al.
(2001:...): K!).
Cardamine pratensis Linnaeus subsp. chinensis O. E. Schulz,
Herbs, perennial, slender, glabrous except for leaves. Rhizomes slender, stoloniferous. Stems
5–25 cm, simple, branched above, flexuous. Basal leaves not rosulate, often withered by
anthesis; petiole 1–1.5 cm; terminal leaflet orbicular, 2–4 mm in diam., with a petiolule 1–4 mm,
sparsely pilose, base rounded, margin entire or obscurely lobed; lateral lobes similar to terminal
one. Middle and upper cauline leaves pinnatisect, margin scabrous with trichomes to 0.1 mm;
petiole 3–8 mm, not auriculate at base; terminal lobe filiform, 1–2.5 cm × 0.4–0.7 mm, base
attenuate and decurrent with adjacent lateral lobes, margin entire, apex acute, not mucronate;
lateral lobes 1–3 pairs, sessile, decurrent, similar to terminal lobe but smaller. Racemes
ebracteate; 2–8-flowered, rachis strongly flexuous; pedicels of young fruit divaricate, 8–14 mm,
slender, soon recurved, glabrous. Sepals ovate, 1.5–2 × 0.8–1 mm, glabrous, membranous at
margin and apex; petals white, obovate or broadly spatulate, 5–6 × 2.5–3 mm; median filament
pairs 2.5–3 mm, filiform; lateral pair 1.5–2 mm; anthers oblong, 0.5–0.6 mm; ovules 8–12 per
ovary. Fruit linear, 1.5–2.2 cm × ca. 1 mm; valves glabrous; style 1–1.5 mm. Seeds pale brown,
oblong, ca. 1.8 × 0.8 mm, wingless.
Distribution: China (Shaanxi, Sichuan).
102
Cardamine subcarnosa (J. D. Hooker) Allan, Fl. New Zeal. 1: 184. 1961; C. hirsuta Linnaeus
var. subcarnosa J. D. Hooker, Fl. Antarct. 1: 5. 1844; C. glacialis (G. Forster) de Candolle
var. subcarnosa (J. D. Hooker) O. E. Schulz, Bot. Jahrb. Syst. 32: 542. 1903. TYPE: New
Zealad, Campbell Islands, Antarctic Expedition 1839–1843, J. D. Hooker s.n. (lectotype
designated by Heenan 2008: 564), BM!).
Herbs, perennial, cespitose. Stems erectto 30 cm. Basal leaves rosulate, pinnately compound,
5–7-foliolate, to 10(–14) cm; petiole 2–4(6) cm, ciliate; leaflets rarely overlapping towards leaf
apex, entire to shallowly toothed, petiolule to 10 mm or rarely absent; terminal leaflet 5–25 × 5–
20 mm, orbicular to broadly elliptic; lateral leaflets 3–2.2 × 2.5–12 mm, elliptic to obovate;
cauline leaves similar to basal but progressively smaller upwards. Racemes bracteate throughout
or basally, elongated in fruit; fruiting pedicels 2–12 mm, ascending to spreading. Sepals 1.3–2.4
mm, oblong, glabrous; petals white or pink to purple, 2.2–4.7 × 0.6–1.3 mm, obovate, apex
obtuse to rounded; stamens 6, filaments 1.5–2.5 mm; anthers 0.3–0.4 mm; ovules – per ovary.
Fruits 9–20 × 0.9–1.3 mm, ascending to spreading; style 1–1.4 mm. Seeds 0.8–1.4 mm, 0.8–1.4
× 0.5–0.9 mm.
Flowering: Nov–Dec.
Habitat: damp places on scree, rock crevices, grassland, peat, fellfields and rushlands in tundra
zone.
Distribution: New Zealand (Campbell Island).
Cardamine tanakae Franchet & Savatier ex Maximowicz, Bull. Acad. Imp. Sci. Saint
Pétersbourg 18: 280. 1873. TYPE:
Cardamine chelidonioides S. Moore, J. Bot. 16: 130. 1878.
Nasturtium tenue Miquel, Ann. Mus. Bot. Lugduno-Batavi 2: 71. 1866; Cardamine tenuis
(Miquel) Koidzumi, Fl. Symb. Orient.-Asiat. 13. 1930. TYPE:
Herbs, biennial herbs, subglabrous or pubescent. Stems 6–25 cm, erect, often branched.
Cauline leaves (lower) 1–6 cm, simple or with 1–3 lateral leaflets on each side of rachis; blade
or terminal leaflet elliptic to orbicular or subreniform, 0.7–3 cm, 0.5–4 cm wide, base obtuse to
subcordate, margin crenate, apex rounded; lateral leaflets ovate to suborbicular, 0.5–2 cm, 0.3–
1.5 cm wide, base obtuse to subtruncate, margin crenate, apex rounded to obtuse; upper cauline
leaves 2–14 cm, with 1–3 lateral lobes on each side of rachis, rarely simple; petiole auriculateat
base; terminal leaflet lanceolate, ovate to orbicular, 0.6–4.2 cm, 0.3–3 cm wide, base cuneate to
obtuse, margin crenate, apex rounded; lateral leaflets elliptic, oblong, or ovate, 0.5–2.2 cm, 0.3–
1.6 cm wide, base cuneate to obtuse, margin crenate or rarely entire. Racemes ebracteate, 0.5–3.5
cm, densely pubescent or rarely subglabrous; fruiting pedicels 5–10(–12) mm. Sepals 2.5–3 mm,
1–1.5 mm wide; petals white (4–)5–6 mm, 2.5–3 mm wide; filaments 2.5–4 mm. Fruits linear,
1.5–2.7(–3.5) cm, 1–1.5 mm wide, often densely pubescent; style 1–2 mm. Seeds 1.2–1.5 mm.
Flowering: Apr–May.
Habitat: forests.
Elevation: to 1200 m.
Distribution: Japan (Honshu, Hyushu, Shikoku).
Cardamine tangutorum O. E. Schulz, Bot. Jahrb. Syst. 32: 360. 1903.
Herbs, perennial. Rhizomes creeping, sparsely scaly, not stoloniferous. Stems (8–)15–30(–
40) cm simple, erect, strongly curved at base, glabrous throughout or sparsely pilose distally.
Rhizomal leaves 7–16(–22) cm; petiole 3.5–8(–12) cm; terminal leaflet oblong, oblonglanceolate, or elliptic, (2–)3–5(–6) × 0.7–1.2(–2) cm, sessile, base cuneate, margin serrate and
ciliolate, apex acute to subobtuse; lateral leaflets 3–5(or 6) pairs, not decurrent at base; Cauline
103
leaves 1–3(–5), (3–)5–10(–13) cm, subleathery; petiole (0.5–)1–2.5(–4) cm, not auriculate at
base; terminal leaflet oblong, 2–4 cm × 3–10(–15) mm, subsessile, glabrous or rarely sparsely
pilose, base cuneate, margin ciliolate and serrate, rarely subentire or double serrate, apex acute or
acuminate; lateral leaflets 2–4 pairs, similar to terminal but slightly smaller, not decurrent at
base. Racemes ebracteate, 10–15-flowered; fruiting pedicels ascending or divaricate, (0.7–)1–1.5
cm, straight, slender. Sepals oblong, 4–5 × (1–)1.5–2 mm, base of lateral pair saccate; petals
purple, spatulate, 0.8–1.4 cm × (2–)3–5 mm, base not clawed, apex rounded; median filament
pairs 6–8 mm, lateral pair 5–6 mm; anthers oblong, 1.2–2 mm; ovules 10–14 per ovary. Fruit
linear, 3.5–4.5 cm × 2–3 mm; gynophore to 2.5 mm; valves smooth, glabrous; style 2–4 mm.
Seeds brown, ovate or suborbicular, 2–2.8 × 1.5–1.8 mm. 2n = 42.
Flowring: May–Jul.
Habitat: montane ditches, swampy meadows, river basins, woodlands.
Elevation: 1300–4400 m.
Distribution: China (Gansu, Hebei, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan).
Cardamine tenera
Cardamine tenuifolia W. J. Hooker, J. Bot. London 1: 247. 1834. TYPE: Australia, Tasmania,
marshes at Formosa, R. W. Lawrence 237 (holotype, K!).
Herbs, perennial. Rhizomes or stolons present, not fleshy. Stems weak, to 1 m. Basal leaves
soon withered; cauline leaves pinnately compound, with 1–5 pairs of lateral leaflets, petiolate,
not auriculate at base; terminal leaflet orbicular to ovate; lateral leaflets ovate to linear of
filifrom, reduced in size upwards. Racemes ebracteate; fruiting pedicels horizontal to spreading,
1–3 cm. Sepals oblong, 2–5 mm; petals white to lilac, 5–12 mm; stamens 6. Fruits linear, 1.5–4
cm × 1–1.5 mm, erect to spreading, rarely overtopping raceme; style 1.5–3 mm. Seeds ovoid to
oblong, 1–2 mm, flattened, with or without a narrow wing.
Flowering: Dec–Jan.
Habitat: swamps, marshy areas.
Distribution: Austalia (New South Wales, SE South Australia, Tasmania, Victoria).
Cardamine tenuirostris W. J. Hooker & Arnott, Bot. Beechey’s Voy. 6. 1830. TYPE: Chile,
Concepcion, collector? (holotype, K).
Cardamine affinis W. J. Hooker & Arnott, Bot. Miscell. 3: 137. 1833; C. tenuirostris subsp.
affinis (Hooker & Arnott) O. E. Schulz, Bot. Jahrb. Syst. 32: 494. 1903. TYPE: Chile,
Valparaiso, Bridges s.n. (holotype, E!).
Cardamine pentaphylla Philippi, Linnaea 33: 6. 1864, non C. pentaphylla W. T. Aiton, Hort.
Kew. ed. II, 4: 101. 1812. TYPE: Chile, Valdivia, Lago Ranco, Jan 1860, Philippi s.n.
(holotype, SGO-49337!).
Cardamine flavescens Philippi, Anales Univ. Chile 41: 667. 1872. TYPE: Chile, mas arriba de
San Fernando, Sep 1864, F. Philippi 63 (lectotype, here designated, SGO-63909!, fragments,
BAA!; isotype, B!).
Nasturtium stenophyllum F. Philippi, Anales Univ. Chile 81: 177. 1893, non N. stenophyllum
Nyman, Consp. Fl. Eur. Suppl. 2, 1: 23. 1889. TYPE: Chile, Valdivia, Futaco, Oct 1865, F.
Philippi 70 (holotype, SGO-45111!; isotype, B!)
Cardamine tenuirostris subsp. reicheana O. E. Schulz, Bot. Jahrb. Syst. 32: 495. 1903. TYPE:
Chile, Ñuble, Oct 1878, Puge s.n. (lectotype, here designated, SGO-63911!).
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Herbs, perennial. Rhizomes slender and sometimes swollen at nodes, not scaly, stem base not
tuberous. Stems (10−)20−55(−75) cm, usually solitary, erect or decumbent, simple or branched
above, glabrous or rarely pubescent above; lowermost nodes often with adventitious roots. Basal
leaves and lowermost cauline compound, 3−9(−12) cm, with 5−17 leaflets; petiole 2−4(−7) cm;
lateral leaflets oblong-linear, linear-lanceolate, linear or filiform, 0.8−2.5 cm × (2−)4−10(−20)
mm, sessile or short petiolulate, entire or shallowly to coarsely dentate, apex acute to obtuse;
upper cauline leaves with 5−15 leaflets, short petiolate to sessile; lateral leaflets linear to filiform,
rarely narrowly oblanceolate, 0.7−3 cm × 0.2−4 mm, sessile, entire or 1−3-toothed, apex acute to
acuminate. Racemes ebracteate; rachis straight; fruiting pedicels (0.5−)0.7−2(−3) cm, divaricate
to ascending, slender, straight. Sepals oblong, (3.2−)3.5−4.5 × 1−1.5 mm; petals white, obovate
to oblanceolate, (7−)8−10(−12) × 2−5(−6) mm, apex obtuse to emarginate, gradually narrowed to
base; filaments 4−6 mm; anthers oblong, 1−1.5 mm; ovules (16−)20−32 per ovary. Fruits linear,
(1.8−)2−4.5(−5) cm × 1.2−2 mm, attenuate to style; style slender, (2−)2.5−5(−5.5) mm. Seeds
brown, ovate, 1−1.8 × 0.8−1.2 mm.
Flowering Oct−Jan.
Habitat: pools, wetlands.
Elevation: 0−1700 m.
Distribution: Argentina (Neuquén, Río Negro), Chile (Región V, VI, VII, VIII, IX, X).
Specimens examined: ARGENTINA. Chubut: Dep. Futaleufú, betw. Trevelin and Corcovado,
Soriano 2954 (BAA). Neuquén: Arroyo Catan-lil, arriba del Fortin, Kurz 6338 (CORD). Dep.
Minas, Dep. Alumine, Lago Norguinco, Rossow et al. 1243 (BAB, BACP); Lago Quillén,
Rossow et al. 1386 (BAB); lags. Epu-Lauquen, Boelcke et al. 10981 (BAA). Dep. Aluminé, lago
Quillén, Balla 3156 (BAA). Dep. Lácar, Parq. Nac. Lanín, Dawson 1355 (BAB). Dep. Los
Lagos, Ea. Fortín Chacabuco, Boelcke & Correa 11738 (BAB). Río Negro. Dep. Bariloche,
Parque Nac. Nahuel Huapi, Arroyo del Jucidio, De Barba 707 (LIL); Bariloche, Meyer 9231
(LIL). CHILE. V. Valparaiso, Bridges s.n. (E).VI. Colchagua, Philippi s.n. (G, W), Bridges s.n.
(K); San Fernando, Philippi s.n. (G, W). VII. Talca, Claude-Joseph 4338 (US); Linares, 2 km N
camine entre Quirihue y Cauquenes, Rodríguez 549 (CONC). VIII. Concepción, San Pedro,
[36°50′S, 73°6′W], Sparre 10003 (CONC); Concepción, Talcahuano, Poeppig 627 (G, W);
Lonquimay, [38°39′S, 70°54′W], 10 Jan. 1947, Pfister s.n. (CONC); Atacalco, [36°53′S,
71°38′W], 30 Nov. 1944, Pfister s.n. (CONC); Candelaria, Los Setenta, [37°22′S, 72°29′W], 2
Nov. 1935, Junge s.n. (CONC); Cordillera de Chillan, 1956−1957, Germain s.n. (G, W). Bíobío:
Termas Los Pemehues-Mulchén, [38°3′S, 71°43′W], 12 Jan 2949, Pinto s.n. (CONC); 15 km N
Cabrero, 37°0′S, 72°21′W, Keeley et al. 25854 (CONC). IX. Temuco, Gunckel 16549 (US),
Claude-Joseph 1007 (US); Cunco, Calude-Joseph 5518 (CONC); Malleco, Termas de Tolhuaca,
[38°14′S, 71°44′W], Zöllner 10303 (MO, US); same area, 12 Dec. 1956, Pfister s.n. (CONC);
between Curacautin and Cherquenco, Smith & Sparre 113HE (CONC, US); Saboya, Montero
5346 (BACP, CONC, SI); Meninco [37°47′S, 72°28′W], Montero 5257 (BACP, CONC); PichiMalleco, Neger s.n. (CONC). X. Futa, Lechler 299 (G, K, P); Panguipulli, Claude-Joseph 2369
(US), Montero 4627 (SI), Montero 6168 (CONC); Osorno, Pucatrihue, [40°32′S, 73°41′W],
Sparre & Smith 288 (CONC); Corral, Isla Manceras, [39°52′S, 73°23′W], 20 Sept. 1932, Junge
s.n. (CONC); Dep. Ancud., Puntra, Sparre 4159 (SGO); Lago Ranco, Montero 3985 (SI).
A highly variable species in leaflet morphology, fruit and style length, and shape and size of
petals. However, none of the taxa treated above as synonyms merit recognition. The
105
characteristic attenuate fruits and long styles easily distinguish the species from the other South
American Cardamine with ebracteate racemes and stems rooting from lowermost nodes.
Except for the collection cited above, I have not yet seen any other material of the species
from Argentina. Those cited by Boelcke & Romanczuk (1984) or annotated by Boelcke in BAA
all represent a mixture of Cardamine variablis, a fewer C. vulgaris, and still fewer C. bonariensis.
For example, Boelcke et al. 14361 is C. bonariensis, Mallo et al. and Correa et al. 7707 are C.
vulgaris, and Boelcke 8994, Boelcke & Correa 11738, Boelcke et al. 13717, and Soriano 2472
are C. vaiabilis. In one collection from Chila, Boelcke 8966 (BAA), most plants have broad
leaflets of basal leaves and tuberous base, but others have narrow leaflets and nontuberous base. I
suspect that C. vulagris and C. tenuirostris are hybridizing, but that needs to be confirmed too. It
is likely that the three species integrade. Many plants have the leaflet form of C. variablis, but for
the number and width of leaflets, they clearly belong to the other two species. Furthermore, C.
vulgaris has tuberous stem base, while C. variablis does not, and one finds rhizomes only slightly
and irregularly swollen. However, these morphological intermediates need to be tested
experimentally and molecularly to see if indeed they represent hybrid derivates.
Cardamine tianqingiae Al-Shehbaz & Boufford, Harvard Pap. Bot. 13: 89. 2008. TYPE: China,
Gansu, Wen Xian: Motianling Shan, Baishui Jiang Nature Reserve, vicinity of town of Fanba,
upstream from Muxüba, 32°41'59"N, 104°53'51"E, 1130–1250 m, remnant mixed deciduous
forest, gravelly river margin, 10 May 2007, David E. Boufford, Qing Tian, Z. Y. Zhang & Y.
Jia 37546(Holotype: PE; Isotypes: A, GAUF, MO).
Herbs, perennial, sparsely pilose on lowermost leaves. Rhizomes thin, 0.5–1 mm wide.
Stems 10–20 cm ascending, branched above, glabrous, flexuous. Rhizomal leaves not rosulate,
pinnately compound, 5–7-foliolate; petiole 3–6 mm, ciliate, pilose adaxially; terminal leaflet 5lobed or toothed, suborbicular to broadly ovate in outline, 3–7 × 3–10 mm, distinctly larger than
lateral leaflets, ciliate, petiolule 3–6 mm; lateral leaflets ovate to subdeltoid, 1–4 × 0.5–3 mm,
entire or obscurely 1- or 2-toothed, ciliate, petiolule 0.1–1 mm; lowermost cauline leaves 5–11foliolate, adaxially sparsely pilose along rachis or glabrous, terminal lobe entire or subapically 3toothed; uppermost cauline leaves 5–7-foliolate; leaflets linear to filiform, 1.5–3.3 cm ×0.5–1
mm, sessile, entire, glabrous, petiole 0.5–1 cm, not auriculate at base. Racemes ebracteate, 5–10flowered; rachis flexuous; flowering pedicels slightly recurved after anthesis; fruiting pedicels
divaricate-ascending, 0.7–1.4 cm, slender, straight, glabrous, not appressed to rachis. Sepals
ovate, 1.5–2 mm, glabrous, subapically membranous, not saccate; petals white, fading pale
lavender, obovate, 4–5 × 2–2.2 mm, flaring, apex obtuse; stamens exserted;median filament pairs
3–4 mm, not flattened; lateral pair 2–2.5 mm; anthers oblong, 0.5–0.6 mm. Fruit 1.3–2
cm × ca. 0.8 mm, ascending; valves glabrous; style 2–3 mm long. Seeds not seen.
Flowering: May.
Habitat: gravelly river margin, remnant sof mixed deciduous forests.
Elevation: 1130–1250 m.
Distribution: China (Gansu).
Notes: known only form the type gathering.
Cardamine trichocarpa A. Richard, Tent. Fl. Abyss. 1: 18. 1847. TYPE: Ethiopia, Begemdir,
Schimper II.1352 (lectotype here designated, P!; isolectotypes, K!, S!, Z).
Cardamine trichocarpa var. elegans Engler, Hochgebirgsfl. Trop. Afr. 225. 1892; C. trichocarpa
subsp. elegans (Engler) O. E. Schulz, Bot. Jahrb. Syst. 32: 463. 1903. TYPE: Ethiopia,
Begemdir/Tigre, Debra-Tabor, Schimper 1162 (holotype, B!; isotypes, BM!, K!, Z).
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Cardamine trichocarpa var. usambarensis Engler, Deutsch-Ostafrika V. Pars C: 183. 1892.
TYPE: not designated.
Cardamine talamontiana Chiov., Ann. Bot. Roma 9: 51. 1911. TYPE: Ethiopia, Semien,
Debarek, Chiovenda 884 (holotype, FI).
Cardamine hirsuta Linnaeus var. subumbellata Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 4:
294. 1852; C. subumbellata (Dalzell) J. D. Hooker & T. Anderson, Fl. Brit. India 1: 138.
1872. TYPE: Belyariss (spelling???) hills, south. (type, E!).
Herbs, annual, subglabrous or substrigose. Stems 5−50 cm, erect, simple or branched from
base. Leaves (all) compound, 1−15 cm, not auriculate at base; lateral leaflets (1 or)2−5 pairs,
oblong to oblong-ovate, 0.5−4.8 cm, 0.4−1.5 cm wide, subsessile or on petiolules to 5.4 mm,
base cuneate or slightly oblique, margin crenate to serrate, apex obtuse to subacute; terminal
leaflet similar to and often larger than lateral ones, on a petiolule 3−12 mm. Racemes ebracteate,
elongated or not in fruit; fruiting pedicels ascending to suberect, 1.4−7 mm. Sepals oblong 0.7−2
mm, sparsely pubescent; petals absent or rudimentary and shorter than sepals; stamens 4, all
median, shorter than sepals; anthers ovate, ca. 0.5 mm; ovules 8−12. Fruits linear, 0.8−2.6 cm,
0.5−1.7 mm wide, erect to ascending, often subumbellate and overtopping flowers; valves
sparsely pubescent or rarely glabrous, attenuate to apex; styles 0.5−1.3 mm. Seeds reddish
brown, broadly oblong, 1.2−1.5 mm, 0.8−1 mm wide.
Habitat: roadsides, clearings, open slightly moist ground.
Elevation: 900−3000 m.
Distribution: Angola, Burundi, Ethiopia, India, Kenya, Sri Lanka, Tanzania, Uganda, Zaire.
Cardamine trifida (Lamarck ex Poiret) B. M. G. Jones, Feddes Repert. Spec. Nov. Regni Veg.
69: 57. 1964; Dentaria trifida Lamarck ex Poiret, Encycl. Méthod. Bot. Suppl. 2: 465. 1811;
Sphaerotorrhiza trifida (Lam. ex Poiret) Khokhrjakov, Fl. Magadan Reg. 235. 1985. TYPE:
Cardamine schulziana Baehni, Candollea 7: 281. 1937. TYPE:
Dentaria tenuifolia Ledeb., Mém. Acad. Imp. Sci. Saint Pétersbourg 5: 547. 1815; Cardamine
tenuifolia (Ledeb.) Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 15: 238. 1842; not W.
J. Hooker, J. Bot. 1: 247. 1834. TYPE:
Dentaria alaunica Golitsin, Del. Sem. Hort. Bot. Univ. Voroneg. 8: 48. 1947. TYPE:
Herbs, perennial. Rhizomes short, with many, slender petioles 0.5–10 cm that terminate into
tubers; tubers fleshy, whitish, representing modified leaf blades, globose, ovoid, or subreniform,
to 7 mm, sometimes slightly flattened and 3–7-toothed. Stems (7–)12–30 cm erect, simple,
glabrous. Rhizomal leaves ternate, biternate, or palmately 5-foliolate; petiole 3–15 cm; terminal
leaflet lanceolate, broadly ovate, or suborbicular in outline, to 2 × 1.3 cm, with a petiolule to 6
mm, base cuneate, margin crenate or dissected into 3 sublanceolate or ovate lobes, base cuneate
or subcordate; cauline leaves 1–3, sometimes 2 and subopposite, often restricted to distal 1/3 of
stem, minutely scabrid at least along margin with trichomes to 0.1 mm, rarely also pilose with
trichomes to 0.4 mm; petiole 0.4–2 cm; terminal leaflet linear or narrowly lanceolate, 0.5–5 cm ×
1–6 mm, base cuneate or attenuate, margin entire or 3-toothed or -lobed, apex acute and
mucronate; lateral leaflets 1 or 2 pairs, similar to terminal one or entire. Racemes 5–20-flowered;
fruiting pedicels ascending, 0.7–1.3 cm. Sepals oblong, 3.5–5 × 1.5–2 mm, glabrous; petals
purple or pink, rarely white, obovate or spatulate, 8–11(–14) × 3.5–5 mm, apex rounded; median
filament pairs 4–6 mm, slender; lateral pair 3.5–4.5 mm; anthers narrowly oblong 1.1–1.5 mm;
107
ovules 10–16 per ovary. Fruit linear, 2–3.5 cm × 1.2–1.6 mm; valves glabrous; style 1.5–4 mm.
Seeds brown, oblong, 1–1.3 × 0.8–1 mm. 2n = 32, 48.
Flowering: May–Jun.
Habitat: moist meadows or slopes, shady places, forests, among shrubs.
Distribution: China (Heilongjiang, Jilin, Nei Mongol), Japan, Kazakhstan, Korea, Mongolia,
Russia (Far East, Siberia).
Cardamine trifolia Linnaeus, Sp. Pl. 2: 654. 1753. TYPE: Herb. Linn. 835.7 (lectotype
designated by Marhold (1996: 127), LINN).
Cardamine trifoliolata J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5: 145. 1861. TYPE:
Bhutan, Griffith 1757 (holotype, K; isotype, BM!).
Cardamine flexuosoides W. T. Wang, Acta Bot. Yunnan. 9: 14. 1977. TYPE: China, Sichuan,
Baoxing, Dengchigou, 3300 m, 26 Jun 1933. T. H. Tu 4291 (wrongly as 21010)(holotype,
PE!).
Cardamine flexuosoides W. T. Wang var. glabricaulis W. T. Wang, Acta Bot. Yunnan. 9: 13.
1987. TYPE: China, Sichuan, Wenchuan, 2500, 28 May 1930, Wang Fatsuan 21010
(holotype, PE!; isotype, KUN!). Same as below.
Loxostemon smithii O. E. Schulz var. wenchuanensis Y. C. Lan & T. Y. Cheo, Bull. Bot. Res.,
Harbin 1(3): 55. 1981. TYPE: China, Sichuan, Wenchuan, 2500, 28 May 1930, Wang
Fatsuan 21010 (holotype, KUN!; isotype, PE!).
Herbs, perennial, often sparsely pilose at least basally. Rhizomes slender, thickened at stem
base, with 1 or few stolons. Stems (4–)6–18(–25) cm, erect or decumbent, slender, simple or few
branched. Basal leaves 1–3, 3- or 5(or 7)-foliolate, rarely simple; petiole 1–4(–5.5) cm; terminal
leaflet broadly obovate or rarely ovate, 2–12 × 3–14 mm, with a petiolule 1–6(–8) mm, base
subtruncate to cordate or rounded, margin entire and obscurely 5-lobed or -crenate, apically
subtruncate or obtusely 3-lobed; lateral leaflets 1(–3) pairs, subsessile or petiolulate, resembling
terminal leaflet, or not lobed and oblong or ovate, smaller; cauline leaves 1 or 2(or 3), 3(or 5)foliolate; petiole 0.5–1.5 cm, not auriculate at base; terminal leaflet similar to that of basal leaf,
with a petiolule 0.5–3 mm; lateral leaflets similar to those of basal leaves. Racemes ebracteate,
lax, 2–8-flowered, rachis straight; fruiting pedicels ascending to divaricate, 0.5–1.5(–2) cm,
straight, glabrous. Sepals ovate to oblong, 2–3 × 1–1.4 mm, glabrous, lateral pair subsaccate;
petals white, obovate to spatulate, 5–8 × 2.5–4 mm, not clawed, apex rounded; median filament
pairs 3–4.5 mm, slender; lateral pair 2–3 mm; anthers oblong, 0.8–1 mm; ovules 8–12 per ovary.
Fruits (young) glabrous. Seeds not seen.
Flowering: May–Jul.
Habitat: moist rocky crevices, meadows, moist forests, mossy banks, rocky areas.
Elevation: 2500–4300 m.
Distribution: Bhutan, China (Sichuan, Yunnan), India (Sikkim), Nepal.
Cardamine tryssa I. Thompson, Muelleria 18: 27. 2003. TYPE: Australia, Victoria, SW corner
of major bridge on Princes Hwy crossing Toorloo Arm of Lake Tyers, between lakes entrance
and Nowa Nowa, 17 Dec 1995, I. R. Thompson 311 (holotype, MEL).
Herbs, annual, glabrous. Stems to 15 cm, erect, slender. Basal leaves rosulate, simple, 2−8
cm; petiole 1−6 cm; blade elliptic to oblong-elliptic or ovate, 0.7−2 x 0.5−1 cm, base cuneate,
margin entire or 1−3 lobes or crenations on each side, apex obtuse to rounded; cauline leaves
0−2, subsessile or petiole to 3 mm, not auriculate at base, obovate to narrowly elliptic, entire or
108
with 1 or 2 crentations on eachs side. Racemes ebracteate, 3−15-flowered; fruiting pedicels
ascending, 5−10 mm. Sepals ovate, 1.3−1.8 mm; petals white inside, pink outside, spatulate,
2.5−4 mm; stamens 6; ovules − per ovary. Fruits linear, erect or nearly so, 2−2.5 cm × 0.7−1
mm; style to 1.5 mm. Seeds elliptic, 0.8−1 mm.
Habitat: open forests near rivers, steep slopes.
Distribution: Australia (Australian Capital Territory, New South Wales, Tasmania, Victoria).
Cardamine tuberosa de Candolle, Syst. Nat. 2: 254. 1821; non C. tuberosa Penzes & Vida,
Ann. Hist. -Nat. Mus. Nat. Hungar., Bot., 57: 174. 1965. TYPE: Chile, Dombey s.n. (holotype,
G-DC; isotypes, B!, 2P!, G!).
Cardamine cognata Steudel, Flora 39: 409. 1856; C. tuberosa subsp. cognata (Steudel) O. E.
Schulz, Bot. Jahrb. Syst. 32: 491. 1903. TYPE: Chile, Rancagua, Bertero 146 (holotype, P!;
isotypes, G!, 2GH!, 2P!).
Cardamine granulata Philippi, Linnaea 33: 8. 1864-1865; C. tuberosa var. granulata (Philippi)
Reiche, Fl. Chile 1: 97. 1896. TYPE: Chile, Andes of Colchagua, Landbeck s.n. (holotype,
SGO, not seen).
Cardamine tuberosa var. velutina Spegazzini, Anal. Mus. Nac. Buenos Aires 7: 212. 1902.
TYPE: Argentina, near Putra-Choique, 1900, N. Illin s.n. (holotype, LP)
Cardamine thyrsoidea O. E. Schulz, Bot. Jahrb. Syst. 32: 591. 1903. TYPE: Chile, Antuco,
Pöppig 1829 (holotype, W!).
Herbs, perennial. Rhizomes tuberous, congested, not scaly, consisting of clusters to 6 × 3 cm.
Stems (8−)15−45(−65) cm, often 1 from tuber, narrowest at base, erect, glabrous or pubescent,
branched above, 3−7-leaved. Basal leaves simple or rarely with trifoliolate; petiole
(2.5−)4−15(−25) cm; leaf blade or terminal leaflet orbicular to reniform, (1.2−)2−5(−7) cm wide,
often wider than long, palmately veined, base often deeply cordate, rarely obtuse, margin crenate
to repand, sometimes distinctly angled and mucronate at vein tips; lateral leaflets (when present)
subsessile or on petiolules to 6 mm, similar in shape and margin and considerably smaller than
termial leaflet. Middle cauline leaves 3−7-foliolate leaflets; terminal leaflet variable in shape,
1−3.5 cm, subentire or dentate; lateral leaflets much smaller; uppermost leaves simple or
compound, smaller and distinctly narrower, often entire. Racemes ebracteate, corymbose,
elongated considerably in fruit; rachis usually straight; fruiting pedicels (0.4−)0.8−2(−3) cm,
ascending to divaricate, straight and often forming a straight line with fruit, often glabrous.
Sepals oblong, (2−)3−4 × 1.2−1.5 mm, erect, caducous, glabrous; petals white, broadly obovate,
8−14 × 4−7 mm, abrubtly narrowed to base to, apex obtuse; filaments 4−6 mm; anthers oblong,
0.7−1 mm. Fruits linear, (1.8−)2−4(−4.5) cm × 1.2−2 mm, attenuate to apex; style slender (1.5)
−2−4(−5) mm. Seeds brown, oblong to broadly ovate, 1.5−2 × 0.8−1 mm.
Flowering: (Jul−)Sep−Dec.
Habitat: among boulders, Nothofagus forests.
Elevation: 0−2700 m.
Distribution: Chile (Región IV, V, Santiago, VI, VII, VIII, IX, X, XI).
Specimens examined: CHILE. IV: Choapa, Cerro Silla del Gobernador, [32°08′S, 71°30′W],
Schlegel 3792 (CONC); Cordillera de Ovalle, La Hualtata, [30°39′S, 70°40′W], Jiles 1224 (M);
Dep. Illapel, Cerro Santa Inés-Centinela-Pichidangui, [31°56′S, 71°22′W], Jiles 4605 (M);
Riecille, Zöllner 6794 (CONC, NA); Ovalle, Río Tascadero-El Polvo, [31°08′S, 70°35′W], Jiles
6382 (CONC). V: El Pangal, Levi 3043 (SGO); Valle El Sauce, near Melón, Zöllner 9352 (MO);
109
Cerro negro, Zöllner 8224 (MO, NY); Cerro La Campana, Hutchison 40 (UC, US), Solbrig et al.
3616 (GH, NY); Cerro El Roble, [32°58′S, 71°00′W], Garaventa 4599 (BAA, CONC, SI); Cajon
de la Vega, [33°03′S, 71°06′W], Garaventa 4612 (BAA, CONC, SI); Viña del Mar, 5 July 1935,
[33°02′S, 71°34′W], Behn s.n. (CONC); Caleu, [33°00′S, 71°00′W], Garaventa 4526 (BAA,
CONC, SI); Laguna Verde, [32°05′S, 71°39′W], 1 Oct. 1922, Behn s.n. (CONC); Valle de
Marga-Marga, Jaffuel & Piron 3050 (GH), Jaffuel s.n. (CONC); Cerro Quebrada Verde,
[33°03′S, 71°42′W], Garaventa 2971 (BAA, CONC, SI); Cajón del Carretón, [32°15′S,
70°56′W], Garaventa 3319 (BAA, CONC, SI); Quilpué, Cerro Buitre, [32°59′S, 71°26′W], 15
Sept. 1935, Behn s.n. (CONC); Cerro Caquicito, [32°45′S, 71°05′W], Schlegel 215 (CONC);
Quillota, Puerta Oca, [32°57′S, 71°05′W], Schlegel 2677 (CONC). Quillota, Bertero 889 (GH,
MO, NY, P); Cerro La Campana, [32°57′S, 71°08′W], Garaventa 2870 (B), Garaventa 2871
(BAA, CONC, SI); El Cricket, [33°03′S, 71°38′W], Garaventa 2226 (BAA, CONC, Si).
Santiago: Quebrada de Macul, [33°30′S, 70°31′W], Pisano & Baraona 1586 (CONC); Cajón del
Maipo, [33°48′S, 70°17′W], Grandjot 4858 (CONC); above jct. rds to La Disputada and
Farellones, [33°20′S, 70°18′W], J. & A. Solomon 4283 (MO, SGO); El Canelo, Cajón del Maipo,
[33°34′S, 70°26′W], Garaventa 2854 (BAA, CONC), Looser 2441 (SI); Cerro Manguehue,
[33°21′S, 70°34′W], Looser 2180 (CONC, SI); Alhué, [33°50′S, 70°56′W], Gunckel 608
(CONC); Mina Las Arañas im Arrayantal bei Los Condes, Grandjot 3208 (CONC, GH);
Santiago, Claude-Joseph 1396 (US), Philippi s.n. (G); Cordillera de Santiago, 1856-1857,
Germain s.n. (G); Río Colorado, Termas del Tupungato, Looser 747 (GH, SI); Tiltil, [33°04′S,
70°58′W], Nov. 1950, Barrientos s.n. (CONC); Valle Macul, [33°30′S, 70°31′W], Sept. 1931,
Grandjot s.n. (CONC, MO, S). VI: Pichilemu, Quebrada El Roble, Villagrán & Pérez 3140
(SGO); Colchagua, Dec. 1869, Philippi s.n. (SGO). VII: Peteroa, Jan. 1933, Grandjot s.n. (MO);
Rauco, [34°55′S, 71°19′W], Barros 2755 (CONC). VIII: Cerros de La Toma, [36°50′S,
73°02′W], 14 Sept. 1935, Junge s.n. (CONC); Cerro Chepe, [36°49′S, 73°04′W], 7 Sept. 1933,
Junge s.n. (CONC), Philippi 1605b (SGO); Atacalco, 18 sept. 1939, Pfister s.n. (CONC); Cerro
Manchou Chico, 8 Sept. 1933, Junge s.n. (CONC); Concepción, Philippi 252 (G); Orillas del rio
Renegado, 2 km de Los Lleuques, [36°52′S, 71°38′W], Rodríguez 199 (CONC). IX: Mininco,
[37°47′S, 72°28′W], 13 Sept. 1952, Schwabe s.n. (CONC); Cordillera de la Costa, above Angol,
Dusén 320a (S); Temuco, Cerro Vielol, [38°43′S, 72°35′W], Montero 7312 (CONC); Fundo
Solano, Los Alpes, Cordiller de Nahuelbuta, Eyerdam 10181 (US). X: Valdivia, 1899, Buchtein
s.n. (US). XI: Aysen, Caleta Vidal, 27 Oct. 1947, Behn s.n. (CONC). Region?: Vichuquen,
Zöllner 4501 (SI); Hualqui, Pichaco, Junge & Zambrano 2056 (SI); Cor de Talca El Picazo,
Barros 2759 (SI)
Schulz (1903) recognizd the laxer and taller form as subsp. cognata, but that distinction is
unwarranted. The species is readily distinguished from the other Chilean Cardamine by having
tuber clusters and stems narrowest at its point of attachment to the tubers. In its overall
morphology, C. cordata is similar to C. rostrata, but the latter has non-tuberous rhizomes and
often angled leaves prominently mucronate at the vein endings.
Cardamine uliginosa Bieberstein, Fl. Taur.-Cauc. 3: 438. 1818. TYPE: Caucasus, Bieberstein
s.n. (holotype, LE).
110
Cardamine uniflora (J. D. Hooker) Allan, Fl. New Zealand 1: 183. 1961; Cardamine hirsuta
Linnaeus var. uniflora J. D. Hooker, Handbook New Zeal. Fl. 12. 1864; C. heterophylla var.
uniflora (J. D. Hooker) Ckn., Rep. Bot. Surv. St. Id. 54. 1909. TYPE: New Zealand, Colenso
1813 (holotype K).
Herbs, perennial?, scapose, glabrous throughout. Rhizomes slender. Stems up to 5 cm, erect,
leafless. Basal leaves rosulate; petiole slender, slightly flattened, to 4 cm; blade simple, ovate to
suborbicular, 0.3–1 cm, entire; cauline leaves absent. Racemes ebracteate 1(–4)-flowered;
fruiting pedicels erect to ascending. Sepals ovate-oblong, ca. 2 mm; petals white, ovate-oblong,
to 7 mm, including claw to 2 mm. Fruits linear, to 2 cm. Seeds < 1 mm.
Habitat: coastal lowlands, montane boggy grounds.
Distribution: New Zealand.
Cardamine umbellata Greene, Pittonia 3: 154. 1897. TYPE:
Cardamine hirsuta Linnaeus subsp. kamtschatica (Regel) O. E. Schulz; C. kamtschatica (Regel)
Piper; C. oligosperma Nutttall var. kamtschatica (Regel) Detling; C. sylvatica Link var.
kamtschatica Regel. TYPE:
Herbs, perennial. Rhizomes often elongated, not fleshy, slender or rarely thickened, 1–2(–5)
mm in diam.; stolons absent. Stems (0.3–)0.8–2.5(–3) dm, erect to ascending, 1 to several from
base, simple, sometimes branched above, not flexuous, glabrous. Basal leaves rosulate, pinnately
compound, (3–)5–7(–9)-foliolate, 2–5(–9) cm, sometime withered at anthesis; terminal leaflet
reniform or orbicular, 0.4–0.8(–1.2) × 0.5–0.9(–1.6) cm, margin entire or 3–(5)-lobed or crenate;
lateral leaflets smaller than terminal, broadly ovate, rarely broadly obovate or orbicular, mostly
entire, rarely slightly 3–(5)-lobed or crenate, shortly petiolulate or subsessile; cauline leaves 3–
5(–7), 3–7(–9)-foliolate; leaflets mostly entire, sometimes 3-lobed or crenate, terminal leaflet
narrowly obovate, ovate, oblanceolate, lanceolate, oblong, lateral ones narrowly obovate,
oblanceolate to linear, subsessile to sessile. Racemes ebracteate, both flowering and fruiting
racemes subumbellate, 2–8(–14)-flowered, rachis usually only 0.3–2 cm; fruiting pedicels 3–8(–
10) mm, suberect to ascending. Sepals oblong, greenish or purplish, 1–2 × 0.5–1 mm; petals
white, narrowly obovate, 2.5–5 × 1–3 mm. Fruit linear, (1.3–)1.8–2.5(–3) cm × 0.8–1.5(–2) mm;
valves glabrous or sparsely pubescent, torulose; style 0.5–2 mm. Seeds brown, oblong, 1–1.5 ×
0.8–1 mm, wingless. 2n = 32, 36, 48.
Flowering: Jun–Sep.
Habitat: streambanks, tundra, alpine slopes, wetlands, damp, swampy and mossy areas, beach
gravel and sand, alpine stream margins.
Elevation: 0–1800 m.
Distribution: Canada (Alberta, British Columbia, Northwet Territories, Yukon), Russia (Far
East/Kamtschtka, Kuriles), United States (Alaska, Washington).
Cardamine variabilis Philippi, Linnaea 33: 5. 1864-1865. TYPE. Chile,Valdivia,Lago Rauco,
1860, Philippi s.n. (SGO-49329!, fragments, BAA!). The type sheet has two collections, of
which the one collected by F. Philippi from Rio Bueno in October 1872 is not the type.
Cardamine ovata Philippi, Anal. Univ. Chile 81: 69. 1892; non C. ovata Bentham, Pl. Hartweg.
158. 1845. TYPE. Chile, valley of río Palena, Jan−Feb 1887, Federico Delfin s.n. (holotype,
SGO-71452!). Although Muñoz (1960) listed SGO-63892! as part of the type collection, this
sheet should not be considered as such because it does not carry Philippi’s hand writing nor
the collector’s name.
Cardamine triphylla Philippi, Anal. Univ. Chile 81: 72. 1892. TYPE: Chile, Valley of río Palena,
Jan−Feb 1887, Federico Delfin s.n. (holotype, SGO-49342!).
111
Cardamine integrifolia Philippi, Anal. Univ. Chile 81: 71. 1892, not C. integrifolia Greene, Bull.
Calif. Acad. 389. 1887, nor C. integrifolia Nasaroff, Bull. Soc. Nat. Mosc. 1923−24, n.s. 32,
341. 1924; Cardamine integrifolia Philippi var. diversifolia O. E. Schulz, Bot. Jahrb. Syst. 32:
431. 1903; Cardamine thermarum Marticorena, Gayana, Bot. 57(2): 191. 2001. TYPE: Chile.
Thermas de Chillán, Feb 1892, Philippi s.n. (holotype SGO-63907!; fragments, BAA!).
Schulz (1903) cited the single collection of the species holotype under his var. diversifolia.
Cardamine integrifolia var. dichondroides Spegazzini, Ann. Mus. Nac. Buenos Aires 7: 212.
1902. TYPE : Argentina, Chubut, prope Lago Blanco, Dec 1900, C. Spegazzini s.n. (holotype,
LP #21135! ).
Cardamine variabilis prol. pinnatisecta O. E. Schulz, Bot. Jahrb. Syst. 32: 432. 1903. TYPE:
Chile, Antuco, 1928, Pöpping s.n. (lectotype, here designated, W!).
Sisymbrium simpsonii Philippi, Anal. Univ. Chile 81: 184. 1892. TYPE: Chile, Río Aisén, E.
Simpson s.n. (holotype, SGO?).
Herbs, perennial. Rhizomes much slender, long, not tuberous or scaly. Stems solitary,
(10−)15−41(−50) cm, erect, 1 to numerous from base, glabrous or sparsely to densely pubescent
throughout, lowermost nodes without adventitious roots. Basal leaves and lowermost cauline
simple and dentate, or compound and 3−7-foliolate, 5−15 cm, glabrous to densely pubescent;
petiole (1−)1.5−6(−7.4) cm; terminal leaflet or blade of simple leaves (1.4−)2−5.8(−7) ×
(0.5−)1−3(−4.2) cm, ovate to obovate or oblong, obtusely to sharply dentate or incised,
sometimes subapically 3-toothed or 3-lobed, rarely repand, petiolule 0.4−1.7 cm; lateral leaflets
ovate to oblong or narrowly oblanceolate to sublinear, 1−3.5 × 0.2−1.5 cm, sessile or on a
petiolule to 5 mm, entire to repand or dentate; upper cauline leaves similar to lower ones in size
or larger, varying from simple to compound, sometimes with l lateral leaflet. Racemes
ebracteate; rachis straight; fruiting pedicels (0.6−)0.8−1.5(−2.2) cm, divaricate-ascending to
horizontal, slender or stout, straight, often forming a distinct angle with fruit. Sepals oblong to
broadly ovate, 3−4 × 1−1.5 mm; petals white, narrowly obovate to oblanceolate, 6−8(−10) × 2−4
mm, gradually narrowed to base, apex obtuse; filaments 4−5 mm; anthers ovate, 0.8−1 mm;
ovules 22−44 per ovary. Fruits linear, (2−)2.5−3.5(−4.7) cm × 1.2−2 mm, attenuate or not at
apex; style slender or rarely stout, 1−2.5(−3) mm. Seeds brown, ovate, 1−1.5 × 0.7−1 mm,
wingless.
Flowering: Oct−Mar.
Elevation: 0−1800 m.
Habitat: rocky areas, wet places.
Specimens examined: ARGENTINA. Chubut: Dep. Cushamen, Lago Puelo, Boelcke et al.
13004 (BAA, BAB, SI), Vallerini 1643 (BAA); Grl. Vintter, Eskuche et al. 1387 (BAA). Dep.
Futalufú, Ea. Pampa Chica, Soriano 2472 (BAA); Tecka, Meyer 9596 (GH, LIL); 22 km W El
Corcovado, arroyo Carbón, Correa et al. 9062 (BAB, BACP); ruta 258, 24 km S del desiro al
lago Rosario, Correa 8995 (BAB, BACP); Trevelin, Estancia Rio Frio, Krapovickas 4006 (BAA,
BAB); entre Trevlin y Corcorado, Ea. Rio Frio, Soriano 2954 (BAA); ruta Nac. 40 y arroyo
Caquel, Correa et al. 9107 (BAA, BAB). Dep. Tehuelches, Lago Vintter, Nicora 9625 (MO, SI),
Nicora 10008 (SI); río Corcorado, Illin 62 (SI), Illin 159 (SI). Neuquén. Hua Hum, Cabrera
11250 (GH). Dep. Catan Lil, Casa de Lata, Cabrera 21895 (SI). Dep. Hiliches, Parq. Nac. Lanín,
Correa et al. 5653 (BAB, BACP); Parque Lanín, San Martín de los Andes, Leal 18164 (BACP).
Dep. Lácar, 32 km SW San Martín de Los Andes, ruta Nac. 234, Fortunato et al. 5768 (BAB);
lago Queñi, Rossow et al. 2088 (BAB). Dep. Los Lagos, Estancia Fortín Chacabuco, Arroyo
Tranquera Colorada, Vallerini 1275 (SI); Los Lagos, Ao Pso de la Cruz, Boelcke 8993 (BAA);
112
Ea. Fortín Chacabuco, Potrero Mula, Boelcke 9030 (BAA); Paso del Córdoba, limits between
Dep. Lácar and Los Lagos, Rossow et al. 2137 (BACP); Los Lagos, Ea. Fortin Chacabuco,
Boelcke & Correa 11738 (BAA); Pto. Manzano, Boelcke et al. 10438 (BAA, BAB). Dep. Minas,
ca. 21 km de Las Ovejas camino a las lagunas Epu-Lauquén, Arroyo de las Bandurrias, Boelcke
et al. 10793 (BAA); between Gendarmería Puerto and Lagunas Epulauquén, Boelcke et al. 11026
(BAA, LIL, MO); Ea. Fortín Chacabuco, Boelcke 8983 (BAA, LIL, MO, SI), Boelcke 8994 (SI);
confluence of rivers Pichi-Nequén and Nequén, cerro de las Yeguas, 36°35′S, 70°45′W, Boelcke
et al. 13717 (BAA, BAB, BACP, SI); Lagunas Epu-Lauqén, Boelcke et al. 10902 (BAA, BAB,
G); Lagunas Epu-Lauquén, Aduana Vieja, 36°50′S, 71°04′W, Boelcke et al. 10821 (BAA, BAB,
LIL, SI). Rio Negro: Dep. Bariloche, San Carlos de Bariloche, 11 Mar 1934, Spegazzini s.n.
(BAA), Buchtein 3 (US); Par. Nac. Nahuel Huapi, Valle de Río Alerce, Boelcke 2084 (SI),
Fabris & Solbrig 670 (LP); Lago Hess, Boelcke 2029 (SI); Pampa Luida, Boelcke & Correa
5670 (BAA, BAB, BACP, SI); Laguna Frias, Boelcke & Correa 5381 (BAA, BAB, SI);
Bariloche, rta 258, río Foyel, Correa et al. 9113 (BAB, BACP). CHILE. VIII: Puerto de
Biobueno, oct. 1871, Philippi s.n. (SGO); Dep. to La Laja, Sierra Velluda, [37°20′S, 71°41′W],
Ricardi 2360 (CONC). IX: Laguna Icalma, [38°47′S, 71°20′W], 13 Jan. 1947, Pfister s.n.
(CONC). X: Riñhué, 1897, Selle s.n. (SGO); Rolecha, [41°55′S, 72°50′W], 5 Jan 1951, Pfister
s.n. (CONC, SI); Pucatrihue, [40°32′S, 73°41′W], Sparre & Smith 289 (CONC), Eyerdam 10538
(UC, US); La Barra del Río Bueno, Sparre 4555 (SGO); Rio Blanco, Andreas 366 (B); Chiloé,
Isla Lagartija, [41°48′S, 72°50′W], Junge 402 (CONC). Valdivia: Llifen-Lago Ronco, [40°19′S,
72°27′W], Montero 3985 (CONC). XI: Lago San Rafael, [46°40′S, 73°50′W], Schlegel 1829
(CONC); Puerto Aysén, Santesson 1236 (S); Aysén Exped., 12 Jan. 1897, Dusén s.n. (S), Dusén
463 (SGO); Puyuhuapi, río Pascua, 10 Oct. 1939, [44°19′S, 72°24′W], Schuabe s.n. (CONC).
Region?: Lago Gullelma, San Carlos de Bariloche, Garaventa 4603 (BACP, SI); Valle Simpson,
16 Dec 1954, Pfister s.n. (CONC).
A highly variable species, as the specific epithet indicates, especially in leaf division, style
length, and number of ovules per ovary. Indeed, a study of the types of the four “species”
included here reveals that the basic differences among them rests solely on leaf shape. The type
of Cardamine triphylla has trifoliolate leaves with narrowly oblanceolate leaflets the terminal one
of which is entire to laterally one-toothed. The type of C. integrifolia is a slender plant with
simple leaves or with a lateral leaflet, and the blad ranges from narrowly obovate to suboblong
and obscurely toothed, whereas the type of C. ovata, annotated by Schulz as C. variablis, also has
simple leaves or laterally with one leaflet, but the leaves are coarsely dentate, and the plant is
stouter. Finally, the type of C. variablis consists of slender plants with 3−5-foliolate lower leaves
the terminal leaflet of which is distinctly larger and strongly toothed but the uppermost leaves
resemble those of types of C. ovata and C. integrifolia in being simple and dentate. On the type
sheet of C. variablis is mounted a second collection with two plants one of which has trifoliolate
leaves and the other with simple leaves. In my opinion, the leaf variation reflected by the types of
these four species represents on a small fraction of what appears to be a complex species with
enormous leaf variation sometimes reflected in the same collection. For example, Santesson
1236 has four plants of which one has pinnatisect leaves with narrowly lanceolate segments,
another with coarsely dentate leaves, and two with strongly incised leaves. It should be admitted,
however, that as delimited here, C. variabilis is not as cleanly defined as the other Chilean and
Argentinian species, and the complex is in need of further studies to elucidate wheter one or
more taxon is involved.
113
I had the chance to examine almost every specimen cited by Boelcke & Romanczuk (1984)
under Cardamine integrifolia. In almost every case, the plants belonged either to C. variabilis or
C. cordate.
Cardamine victoris
Cardamine violacea (D. Don) Wallich ex J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5:
144. 1861; Erysimum violaceum D. Don, Prodr. Fl. Nepal. 202. 1825. TYPE: Nepal, GossainThan, Wallich 4782 (holotype, K; isotype, E!).
Cardamine violacea subsp. bhutanica Grierson. TYPE: Bhutan. Pele La, Greirson & Long 546
(holotype, E!).
Herbs, perennial, shortly pilose or subglabrous. Rhizomes stout, to 1 cm in diam. Stems 20–
100 cm, erect, simple, stout, glabrous. Basal leaves not seen, often withered by anthesis; middle
cauline leaves sessile, strongly auriculate, sagittate, to amplexicaul at base; blade lanceolate to
linear-lanceolate or ovate-lanceolate, 3.5–20 × 0.7–3.5 cm, abaxially glabrous, adaxially
puberulent to subpilose, margin ciliolate and dentate, denticulate, or entire, apex acuminate to
caudate; auricles oblong to ovate, 2–10 × 2–7 mm. Racemes ebracteate, 5–25-flowered;
flowering pedicels spreading to reflexed; fruiting pedicels divaricate to ascending, 0.8–3 cm,
glabrous, straight. Sepals oblong 5–7 × 1.5–3 mm, glabrous or sparsely pilose, base of lateral
pair saccate; petals purple, spatulate to obovate, 1–1.7 cm × 4.5–7 mm; median filament pairs 7–
9 mm, lateral pairs 6–7 mm; anthers oblong, 1.5–2 mm; ovules 10–16 per ovary. Fruit linear, 2–
6 cm × 1.4–2.5 mm; gynophore 0.5–1 mm; valves smooth, glabrous; style 3–8 mm. Seeds
brown, oblong, 2–3 × 1.4–1.8 mm, wingless.
Flowering: May–Aug.
Habitat: grassy slopes, streamsides, open forests, pastures, thickets, roadside banks, sandy moist
forests, forest ravines.
Elevation: 1800–4000 m.
Distribution: Bhutan, China (Yunnan), India (Sikkim), Nepal.
Cardamine volckmannii Philippi, Linnaea 33: 7. 1864-1865. TYPE: Chile. Coquimbo: Baños
del Toro, [29°50′S, 70°02′W], [3260 m], 1860, Volckmann s.n. (holotype, not at SGO
(Muñoz-Schick, pers. com.); isotype, CONC-63192!). No specimens were listed by Muñoz
Pizarro (1960).
Cardamine nivalis Gillies in W. J. Hooker, Bot. Miscell. 3: 136. 1833; C. hirsuta Linnaeus var.
nivalis (Gillies) J. D. Hooker, Fl. Antarct. 3: 232. 1845; non C. nivalis Pallas, Reise Russ.
Reich. 2: 113. 1773. TYPE: Chile, upper spring below the Cumbre, Gillies s.n. (holotype, E!).
Cardamine stricta Philippi, Anal. Univ. Chile 81: 77. 1892. TYPE: Chile, Andes of Prov.
O’Higgins, Cordillera del Peuco, [33°56′S, 70°30′W], 1886, Cádiz s.n. (lectotype, here
designated, SGO-71637!). Another sheet of the type collection, SGO-71638!, carries the
handwriting of F.E. Philippi instead of R. A. Philippi, and it should serve as the isolectotype.
Both sheets lack the lowermost portion of the plant and doubtfully belong to C. volckmanii
because the lateral leaflets are dentate, pubescent, and sessile. Muñoz (1960), listed a third
specimen, SGO-63884!, as part of the type collection under both C. stricta and C. andina, but
that number clearly belongs to the latter taxon, which treated in this account as a synonym of
C. variabilis.
114
Cardamine petiolulata Philippi ex O. E. Schulz, Bot. Jahrb. Syst. 32: 520. 1903. TYPE: Chile,
Cordillera of Santiago, Valle del Yeso, Jan 1866, Philippi s.n. (lectotype, here designated,
SGO-63905!).
Cardamine nivalis subsp. Dep. auperata O. E. Schulz, (check his monograph and citations). GH
has one collection: las Condes, Nov 1871, Reed s.n. (GH!).
Herbs, perennial, glabrous throughout. Rhizomes not tuberous, slender, not scaly. Stems
often single from base, 8−26(−35) cm, erect, glabrous, unbranched above. Basal leaves absent;
lowermost cauline leaves pinnately compound, 2−12(−27) cm, with 2−4(−5) pairs of lateral
leaflets, always glabrous; petiole (1−)2−14(−25) cm; terminal leaflet orbicular to broadly obovate
or ovate, (0.8−)1−2.5(−3) × (0.7−)1−2 cm, base obtuse to cuneate, margin entire or repand,
sometimes obtusely 3-lobed with minutely mucronate tips, apex rounded, petiolule 2−10 mm;
lateral leaflets distinctly smaller or subequalling terminal one, opposite or alternate, 0.5−1.5(−2)
× 0.2−1.4(−1.7) cm, obovate to ovate or suboblong, base cuneate, entire or repand, apex rounded;
uppermost leaves similar to lower ones but smaller and sessile or with petiolutes 1−5(−10) mm.
Racemes ebracteate, corymbose, elongated considerably in fruit; rachis straight; fruiting pedicels
4−10 mm, suberect to ascending, straight, glabrous. Sepals oblong, 2.5−4 × 1−1.5 mm, erect,
caducous, glabrous; petals white, oblong-obovate, 6−9(−12) × 2.5−4(−5) mm, not clawed, apex
obtuse; filaments 3−6 mm; anthers oblong, 0.7−1 mm; ovules 20−34 per ovary. Fruits linear,
(2−)2.5−3.5 cm × 1−1.3 mm, attenuate to apex; style 1−2 mm. Seeds brown, oblong to broadly
ovate, 1−1.5 × 0.8−1 mm.
Flowering: Dec−Mar.
Habitat: wet soil in bogs near streams, very wet stream banks, seepy places, floodplains.
Elevation: 2000−3800 m.
Distribution: Argentina (La Rioja, Mendoza, Neuquén, San Juan), Chile (Región III, IV,
Santiago).
Specimens examined: ARGENTINA: La Rioja. San Juan, Hosseus 1559 (CORD). Mendoza:
Mt. Aoncagua, Puente del Inca, 21 Dec. 1946, Wall s.n. (A), Malme 2851 (S), Boelcke 8912
(BAA). Dep. Las Heras, Punete del Inca, Sparre 1558 (LIL, S), Hauman 116 (G), F. Philippi 61
(SGO); Puente del Inca, Banderita Norte, Boelcke et al. 9787 (BAA, BAB, SI), Boelcke 9809
(BAA, BAB, SI); Las Cuevas, Quebrada de Matienzo, Wingenroth et al. 88 (SI); Tunuyán,
arroyo de la cascada La Vieja, Boelcke et al. 10114 (BAA, BAB, SI); Tupungate, Estancia La
Carrera a Tupungate, Melis & Paci 9 (GH). Neuquén: Minas, Cajón d los Chenques, 26°28′S,
70°48′W, Boelcke et al. 13963 (BAA, BACP, SI). San Juan: Iglesia, Quebrada del Agua Negra,
Cabrera et al. 27065 (BAA, BAB, LP, MO), Cabrera et al. 31147 (SI), Kiesling 6729 (NY, SI).
Dep. Calingasta, El Pachon, de casa Amarilla a Casilla Naranja, Kiesling & Sáenz 1413 (SI); Los
Patillos, Kurtz 9671 (CORD). Dep. Calingasta, Sapitos, Valle Santa Cruz, Luti 10003 (SI); entre
“El Molle” y “Ojo de Agua,” Luti et al. 5575 (SI). CHILE. Los Andes, Potrero Escondido,
Boelcke 2459 (BAA). Valle del Río Blanck, Boelcke 2434 (BAA) III: Vallenar, Cordillera
Laguna Chica, [29°51′S, 69°42′W], Werdermann 257 (CAS, CONC, E, F, G, GH, LIL, MO, S,
SI, UC); vicinity of Laguna Chica, 28°48′S, 69°52′W, Johnston 5968 (GH); Rio Sancarron at
Corrales, Johnston 6212 (GH); Laguna Grande, 28°43′S, 69°56′W, Johnston 5920 (GH);
Quebrada Los Barriales, [29°16′S, 70°09′W], Arancio et al. 94025 (CONC). IV: Cordillera Doña
ana, Cancha de Sky, [28°48′S, 70°03′W], Squeo & Marticorena 88163 (MO); Dep. Elqui,
camino San Juan, Lado sur del Embalse La Laguna, [30°38′S, 70°17′W], Ricardi et al. 1753
115
(CONC); Borde del Cerro Tapado, [30°12′S, 70°02′W], Arroyo 81089A (CONC); Limari,
Cordillera de Ovalle, Río Gordito, [31°02′S, 70°20′W], Jiles 2542 (CONC); Cordillera de
Ovalle, Quebrada Calabozos, Jiles 2940 (CONC); Cord. Río Tascadero, [31°05′S, 70°35′W],
Jiles 6430 (CONC); Bocatoma, Los Molles, Ruthsatz 6083 (MO); Dep. to Illapel, La Vega
Escondida, E Cuncumen, Morrison 16554 (DS), Morrison 16954 (DS, UC, US); Cerro
Curimahuida, 10 km E Matancilla, Worth & Morrison 16676 (DS, G, UC, US); Illapel, Philippi
1605a (SGO); Valle del Choapa, Hacienda de Cuncumen, Looser 2407 (SI). Dep. Elqui, La
Laguna, 25 Apr 1947, Wagenknecht s.n. (SI). Elqui. Canchas de Sky, 29°51′S, 70°3′W, Squeo
88163 (CONC). V: Dep. La Ligua, Junta de Piuquenes, Morrison 17315 (DS, UC, US);
Caracoles, Pennell 13006 (F, GH, NY, SGO); Los Maitenes near Río Colorado, [32°42′S,
70°25′W], Zöllner 6579 (CONC, MO), Zöllner 4388 (SI); Quebrada al fondo de Lag. del Inca,
[32°50′S, 70°08′W], Ricardi 2974 (CONC); Valle Juncal, [32°53′S, 70°10′W], Schlegel 1476
(CONC), Zöllner 9145 (CONC). Santiago: 2.5 km from Villa Paulina, Sweeney 270 (MO); Lo
Valdéa, [33°50′S, 70°05′W], Jan 1950, Barros s.n. (CONC), Gunckel 57035 (CONC); Valdés
tál, Jan 1936, Grandjot s.n. (MO); Valle de Maipo, [33°50′S, 70°05′W], Garaventa 6284
(BACP, CONC, SI), Ruthsatz 6815 (MO); Cord. Vale Morales, 30 Dec. 1940, [33°50′S,
70°05′W], Schwabe s.n. (CONC); Los Condes, Fierro Carrera, [33°12′S, 70°14′W], Garaventa
551 (BAA, BACP, CONC, SI), Garaventa 1638 (BAA, BACP); between Disputada and Perez
Caldera, [33°10′S, 71°18′W], Sparre 11038 (CONC); Potrero Grande, Zöllner 4620 (SI); Fierro
Carrera, Garaventa 1714 (SI), Garaventa 1717 (SI); S. José de Maipo, Cajón del río Morales,
33°30′S, 70°0′W, Saavedra & Pauchard 80 (CONC); San José de Maipo, Cajón del Morales,
33°46′S, 70°40′W, Teillier & Márquez 5332 (CONC); Farellones, Böcher et al. 664 (BAA).
Schulz (1903) adopted the later homonym Cardamine nivalis as a legitimate name apparently
because the earlier homonym has been considered as Macropodium nivale (Pallas) R.Br. since its
tansfer by Brown in 1812. However, that handling of the nomenclature is erroneous, and the
earliest name for the present species is C. volckmanii.
Schulz also listed Cardamine glacialis (G. Forster) DC. var. elatior A. Gray in the synonymy of
C. nivalis, but the latter, recognized herein as ….., does not occur in Tierra del Fueguo, where the
type of var. elatior was collected, nor does C. volckmannii, a species that Schulz reduced to
synonymy of C. glacialis.
Morrison 16954 is a mixed collection of Cardamine volckmannii and C. bonariensis.
Cardamine vulgaris Philippi, Linnaea 28: 665. 1856; C. hirsuta Linnaeus var. vulgaris
(Philippi) Reiche, Fl. Chile 1: 100. 1896. TYPE: Chile, Valdivia, Philippi s.n. (lectotype,
designated by Muñoz (1960), SGO-49363!).
Cardamine intermedia Steudel, Flora 39: 410. 1856; non C. intermedia Hornemann, Fors.
Dansk. Oecon. Pl 3, ed. 1, 714. 1821; nec C. intermedia W. J. Hooker, Icon. Pl. 3: 258. 1840.
TYPE: Chile, Valparaiso, Aug 1829, Bertero 1793 (holotype, P!; isotypes, 3G!, GH!, NY!).
Cardamine macrostachya Philippi, Anal. Univ. Chile 81: 75. 1892. TYPE: Chile, Araucania,
Nov 1887, Philippi s.n. (holotype, SGO-63906!). Muñoz (1960) also listed SGO-49420
(fragments, BAA!) as part of the type collection, but I have not seen that.
Cardamine vulgaris Philippi var. oligozyga O. E. Schulz, Bot. Jahrb. Syst. 32: 544. 1903. TYPE:
Chile, Araucania, Nov 1887, Philippi s.n. (lectotype, here designated, SGO!).
Cardamine grandijotii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 12: 39. 1934. TYPE:
Chile, Cordillera of Santiago, Potrero grande, am sumpfigen Bach, 2200 m, 9 Dec 1933, C.
Grandijot 1 (holotype, B!).
116
Cardamine garaventae O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem: 12: 208. 1934. TYPE:
Chile, Cerro de la Campana, cerca del lugar denominado “La Gotera,” 1600 m, 2 Oct 1932,
Augustin Garaventa 2872 (holotype, B!; isotypes, BAA!, BACP, 2SI!).
Herbs, perennial. Rhizomes much thickened, short, forming a tuberous stem base, not scaly.
Stems solitary, 10−45(−60) cm, erect, simple or branched above, glabrous or sparsely to densely
pubescent, lowermost nodes without adventitious roots. Basal leaves and lowermost cauline
compound, 2.5−20 cm, with (3−)5−11(−19) leaflets; petiole (1−)2−8(−11) cm; terminal leaflet
(0.5−)1−2.5(−3.5) × 0.9−3(−3.5) cm, ovate to obovate or suborbicular or subreniform, dentate to
subcrenate or repand, rarely entire, often petiolule 2−13(−20) mm; lateral leaflets similar or
broadly ovate to oblong, 4−12(−17) × 1−7(−10) mm, sessile or on a petiolule 1−5(−10) mm,
entire or shallowly to coarsely dentate; upper cauline leaves similar to lower ones but smaller and
with sessile or subsessile, narrowly ovate or oblong to linear leaflets. Racemes ebracteate; rachis
straight; fruiting pedicels (4−)6−15(−20) mm, divaricate to ascending, slender, straight, forming
a distinct angle with fruit. Sepals oblong, 2.5−3.5 × 1−1.5 mm; petals white, narrowly obovate to
oblanceolate, (5−)6−9 × 2−3.5 mm, gradually narrowed to base, apex obtuse; filaments 3.5−5
mm; anthers oblong, 0.9−1.1 mm; ovules 20−34 per ovary. Fruits linear, 2−3.5(−4) cm × 1−1.5
mm, attenuate to style; style slender, (1.5−)2−3(−3.5) mm. Seeds brown, ovate, 1.2−1.6 ×
0.8−1.2 mm, wingless.
Flowering: Oct−Mar.
Elevation: 0−3400 m.
Habitat: banks of ditches, freshwater marsh, shrubby areas, disturbed ground in moist
Northofagus forest, wet areas among boulders.
Distribution: Argentina (Chubut, Neuquén, Río Negro, Tucumán), Chile (Región V, VI, VII, IX,
X, XI, Juan Fernandez Is.).
Specimens examined: ARGENTINA. Lago Machónico, Garaventa 4608 (SI). Chubut: Curatel
Epujín, Lourteig & Buchinger 47 (P). Dep. Futaleufú, lago Futaleufqúen, Correa et al. 4168
(BACP); Lago Rivadavia, Correa et al. 8960 (BAA, BAB); Río Tecka, Ea. Pampa Chica,
Soriano 3819 (BAA). Dep. to Rio Senguirr, Lago Blanco, 4 Dec 1903, Koslowsky s.n. (NA);
Lago La Plata, Soriano 3150 (BAA). Dep. Tehuelches, Lago Vintter, Nicora 7420 (SI), Nicora
9477 (SI); Río Corcorado, Illin 132 (SI, UC). Neuquén: Dep. Huiliches, Hosteria Lolog, Boelcke
et al. 6646 (BAA); Parq Nac Lanín, volcan Lanín, Correa et al. 5597 (BAA, BAB); Lago
Quillén, 31 Jan 1963, Valla et al. s.n. (BAA); between Pichi Traful and Lago Villarino, Correa
et al. 7797 (BAB, BACP); Va. La Angostura, area alberque Inacayal, orilla Nahuel Huapi, 27 Jan
1982, Mallo et al. 18172 (BAA, LIL, MO); Lago Correntoso, Cabrera 5070 (F); Lago Aluminé,
Correa et al. 8114 (BAA, BAB, BACP); Lago Quñi, Gentili 810 (SI). Dep. Catan Lil, Paso de
Rahue Vega, Cabrera et al. 32982 (SI); Puerto Manzano, 4 Feb. 1967, Diem s.n. (SI). Dep.
Lácar, RP 23, camino a los baños del Quueñi, Correa et al. 9360 (BAA, BAB). Dep. Los Lagos,
Ea. Fortin Chacabuco, Mallin Medio, Boelcke & Correa 11741 (BAA); Ea. Fortín Chacabuco,
Boelcke 8988 (BAA, G, MO, SI); ruta J, 0.5 km N del arroyo Cla-Clo, Rossow & Gómez 412
(BAA); Parq. Nac. Nahuel Huapi, Ea. Fortiu Chacabuco, Boelcke & Hunziker 3571 (BAA); Parq.
Nac. Nahuel Huapi, Isla Victoria, Boelcke 1794 (BAA, SI). Dep. Picunches, Pino achado, Puest
Gendarmería, 24 Jan 1963, Valla et al. s.n. (BAA); Cerro Otto, Boelcke & Hunziker 3706
(BAA). Río Negro: Dep. Bariloche, Bariloche, Hermann 3469 (NY, SI); El Bolsón, Rasp 13
(SI); Los Cantaros, Boelcke & Correa 5269 (BAB, SI); Valle del Río Alerce, Boelcke 2048
(BAA); Lago Guiterrez, Boelcke & Hunziker 3332 (BAA); Pto Manzano, Boelcke & Correa
11734 (BAA). Tucumán: Dep. Chiclgasta, arroyo Celeste, R 65, km 28, 27°20′S, 65°50′W,
117
Boelcke et al. 5465 (BAA), Boelcke et al. 5464 (BAA); Estancia Las Povas, Venturi 4618 (CAS,
GH, LIL, LP, MO, NY, SI). CHILE. Fundo Trafún, orillas Río Reyhueico, Boelcke 8966 (BAA);
this collection is cited in the text of C. tenuirostris. V: Valparaiso, Claude-Joseph 3629 (US);
Cerro de la Campana, [32°57′S, 71°08′W], Nov. 1950, Collantes s.n. (CONC), Garaventa 3054
(BACP, CONC, SI), Hutchison 40 (GH); San Felipe, Cerro Tabaco, [32°39′S, 70°49′W],
Schlegel 813 (CONC); Quebrada del Cricket, [33°03′S, 71°38′W], 4 Oct 1936, Behn s.n.
(CONC), Garaventa 8272 (SI); Cerro la Campana, Trayecto Placa Darwin a Mina, Villagrán &
Meza 3182 (SGO). Santiago: Farellons, Villagrán & Meza 370 (SGO); Potrero Grande, Zöllner
4621 (SI); Volcán San José, Feb. 1950, [33°47′S, 69°55′W], Castillo s.n. (CONC); Potrero
Grande, Jan 1936, Grandjot s.n. (MO, SI); San Gabriel, Valle Maipo, [33°26′S, 70°14′W],
Schlegel 4409 (CONC). VI: San Fernando, [24°56′S, 70°25′W], Montero 7391 (CONC). VII:
Constitución, [35°20′S, 72°25′W], Barnier 109 (CONC). VIII. Concepción, Gunckel 671
(CONC); Camino de Concepción a Copiulemu, Fdo. San Nicolás, [36°54′S, 72°44′W], Montero
& Montero 34 (CONC); Atacalco, 35 Nov. 1944, [36°53′S, 71°38′W], Pfister s.n. (CONC);
Santa Barbara, Puente Mininco, 1 Nov. 1943, [37°40′S, 71°59′W], Pfister s.n. (CONC); ca. 10
km S rd to Curanilahue along rte 160, 37°30′S, 73°30′W, Taylor 10595 (CONC, MO); San
Pedro, Sparre 9943 (CONC); Talcahuano, San Vicente, [36°46′S, 73°08′W], Sparre 9952
(CONC), Philippi s.n. (SGO); E Chillán from Refugio El Aserradero on rd to Termas de Chillán,
36°54′S, 71°27′W], Taylor et al. 10247 (MO); Mulchén, [37°44′S, 72°14′W], Garaventa 4617
(BAA, CONC, SI); Florida, Fundo Colico, 20 Sept. 1977, [36°49′S, 72°43′W], Oehrens s.n.
(CONC); Termas de Chillán, 36°50′S, 71°25′W, Taylor & Taylor 10847 (CONC, MO), road
from Hualpencillo to Museo de Hualpen, [36°47′S, 73°08′W, Lammers et al. 7511 (CONC,
MO); Chillán, Bureo, Garaventa 2788 (SI). IX: Isla Moscha, [38°22′S, 72°55′W], Gunckel
33265 (CONC); Puerto Saavedra, [38°47′S, 73°23′W], Hollermayer 173 (CONC); Lumaco
(Santa Clara), [38°12′S, 72°49′W], Kunkel 543 (CONC); Pucon, Caburga, [39°12′S, 71°49′W],
Montero 9313 (CONC); Contulmo, [37°59′S, 73°15′W], Nov. 1942, Gunckel s.n. (CONC);
Santa Barbara, Lomas a la izquierda del Pte. Mininco, [37°40′S, 71°55′W], Pfister 680 (CONC);
Reserva Forestal Malleco, Orillas del río Niblinto, [39°10′S, 71°45′W], Marticorena & Quezada
1537 (CONC); Reserva Forestal Huerquehue, [39°10′S, 71°42′W], Baeza 137 (CONC);
Pillanlelbún, [38°38′S, 72°27′W], Montero 5378 (CONC); Pass Longuimay, Zöllner 8142 (MO);
Dep. Victoria, road to Banos Calientes, Morrison & Wagnecht 17507 (UC). X. Valdivia, Isal
Teja, Gunckel 16943 (CONC); Rio Palena, Fan-Feb 1887, Delfin s.n. (SGO); San Juan, Sept.
1872, F. Philippi s.n. (SGO); Collico, Santesson 973 (S), Buchtein s.n. (F), Gunckel 2495
(CONC, GH); Cerro Marina, [39°53′S, 73°25′W], Gunckel 1800 (CONC, GH); La Aguada, same
coordinates, Gunckel 1855 (CONC, GH); Llanquihue, Lago Tods los Santos, March 1974,
[41°08′S, 72°25′W], Villagrán s.n. (CONC); Rolecha, [41°55′S, 72°50′W], 5 Jan 1951, Pfister
s.n. (CONC); Dep. to Castro, Lago Huillinco, [42°42′S, 73°50′W], Marticorena et al. 136
(CONC); Cuesta de Soto, [39°47′S, 73°11′W], Gunckel 1801 (CONC, GH); Quitaluto, [39°57′S,
73°27′W], Gunckel 1538 (CONC); Corral, Gunckel 1846 (CONC, GH); Quebrada del Bolsón,
[39°54′S, 73°26′W], Gunckel 2596 (CONC, GH). XI: Aysen, Puyuhuapi, [44°21′S, 72°34′W],
13 Apr. 1939, Schwabe s.n. (CONC); Estancia Ñirehuao, 20-30 km N Rio Coyhaique, Santesson
1312 (S). Juan Fernandez Is.: Masafuera, Skottsberg 479 (B); Quebrada Pasto, Stuessy &
Lammers 9397 (CONC).
118
The type of Cadamine vulgaris is a small plant in flower, and except for its larger flowers, it
can be confused C. marginata (see below). On the other hand, C. macrostachya is a robust form
of C. vulgaris, a species in which plant size is highly variable as judged from the description
above. The type collection of C. intermedia can hardly be different from that of C. macrostachya,
but the name, though is the earliest published for the species, cannot be taken up because it is a
later homonym. The overall aspects of the type collection C. garaventae in flowers and young
fruits is not any different from those of C. macrostachya and C. intermedia, and the major
difference is that the lower leaves have larger and wider leaflets and longer petiolules that those
of the latter two “species.” Finally, the holotype of C. gandjotii is a pubescent form of C.
vulgaris, and the leaves have lager terminal leaflets and sessile lateral ones. It can easily be
concluded that the variation represented by the type collections of the the above taxa is only in
the leaves, and a comparison of the flowers and fruits clearly support the placement of these taxa
in one species. In fact, the overall variation of the species in leaflet morphology is much higher
that what is represented by those in the types of the above synonyms.
Schulz (1903) treated Cardamine macrostachya and C. vulgaris as inDep. endent species, but
a comparison of their types show that the former is robust and the latter is slender plant. In their
overall aspects of morphology, the two cannot be maintained as distinct taxa, especially if a large
series of plants are critically examined. He also associated C. macrostachya with C. tuberosa and
C. vulgaris with C. glacialis. However, in C. tuberosa the tubers are clustered, whereas in C.
vulgaris (including C. macrostachya) and C. glacialis only the stem base is tuberous, though
both sometimes do not produce the typically tuberous base. Cardamine vulgaris can easily be
distinguished from C. glacials by having attenuate fruits with slender styles, divaricate to
divaricate-ascending fruiting pedicels that often form a distinct angle at their attachment with
fruit, and basal and cauline leaves drastically different in leaflets width. In C. glacialis the styles
are blund, the fruits and fruiting pedicels are often appressed to stem and predominantly forming
a straight line or so, and cauline and basal leaflets not drastically different morphologically.
Because of their slender styles, narrowly oblong to linear or filiform uppermost leaflets, and
divaricate or divaricate-ascending fruiting pedicels, Cardamine vulgaris can be confused with C.
tenuirostris. However, C. vulgaris has tuberous stem base, short rhizomes, ovate to broadly
oblong leaflets of basal and lowermost caulineleaves, fruiting pedicels forming an angle with
stem, and styles (1.5−)2−3(−3.5) mm long. Cardamine tenuirostris often has slender rhizomes
very rarely swollen at nodes, oblong-linear, lanceolate-linear, linear to filiform leaflets of basal
and lowermost cauline leaves, fruiting pedicels forming a straight line with fruits, and styles
(2−)2.5−5(−5.5) mm long.
Cardamine vulgaris is highly variable in leaf divisions, and forms with the broadest leaflets
were described by Schulz (1934) as C. garaventae. Although this form occupies the
northernmost range of the species in Región V and its leaflets of upper leaves are not much
narrower than those of the lower and basal leaves, it is indistinguishable from typical plants of C.
variablis especially in flowers, fruits, styles, ovule number, and presence of tuberous stem base.
Schulz divided this species and the others (e.g., C. glacialis and C. bonariensis), which are highly
variable in leaf morphology, into several infraspecific taxa (e.g., subspecies, proles, varieties,
forms) and species regardless to their similarities in other structure. In my opinion, in the absence
of detailed systematic studies, this fine splitting of species is unwarranted.
Cardamine waldsteinii Dyer, Kew Handlist Herb. Pl. 97. 1891; Dentaria trifolia Waldstein &
Kitaibel, Descr. Icon. Pl. Hung. 2: 148. 1805. TYPE: “in subalpinis Croatiae,” (lectotype
119
designated by Chrtek & Skočdopolová (Sborn. Nár. Mus. Praze, Řada B, Přír Vědy 38B: 222.
1982), PR no. 155765/750).
Cardamine yezoensis Maximowicz, Bull. Acad. Imp. Sci. Saint Pétersbourg 18: 277. 1873.
TYPE: Japan, Hakodate, Monidzi, May 1861, Maximowicz s.n. (holotype, LE!; isotype, P!).
Cardamine akitensis Mochizuki, J. Jap. Bot. 44: 340. 1969.
Cardamine fauriae Franchet, Bull. Soc. Philon. Paris, Ser. VII. 12: 83 (1888), not C. fauriei H.
Léveillé (1912). TYPE: Japan, Yeso, Mt. Otaru, 27 May 1885, Faurie 183 (lectotype here
desingated, P!; isolectotype, P!).
Cardamine fauriae Franchet var. incisa H. Boissieu, Bull. Herb. Boiss. 7: 783. 1899.
Cardamine fauriae Franchet var. oblonga H. Boissieu, Bull. Herb. Boiss. 7: 783. 1899.
Cardamine fauriei H. Léveillé, Repert. Sp. Nov. Regni Veg. 10: 350. 1912, not C. fauriae
Franchet (1888); C. mariformis Nakai, Bot. Mag. (Tokyo) 26: 324. 1912. TYPE: Korea, secus
torrents montis des diamants, 23 Jun 1906, P. U. Faurie 557 (holotype, E! ; isotype, BM!).
Cardamine geifolia Koidzumi, Ic. Pl. Koiz. 2: t. 97. 1914.
Cardamine kiusiana H. Hara, J. Jap. Bot. 14: 516. 1938.
Cardamine torrentis Nakai, Bot. Mag. (Tokyo) 42: 479. 1928; Cardamine yezoensis
Maximowicz var. torrentis (Nakai) Ohwi, Fl. Jap., ed. 2, 671. 1965.
Cardamine valida (Takeda) Nakai, Bot. Mag. (Tokyo) 42: 26. 1928. (basionym??)
Cardamine yezoensis Maximowicz var. kiusiana (H. Hara) Ohwi, F. Jap., ed. 2, 671. 1965.
Herbs, perennial glabrous or sparsely hairy above, sometimes stoloniferous. Rhizomes
slender or thickened, creeping. Stems (15–)25–60(–80) cm, erect, branched above. Cauline
leaves (3–)6–20 cm; lateral leaflets (1 or)2–4(–6) on each side of rachis, oblong-lanceolate,
elliptic, ovate to suborbicular, (0.5–02–3(–3.5) cm, (0.3–)1.5–2.5 cm wide, base obtuse to
subcordate, margin subentire or coarsely few toothed, apex obtuse to rounded; petiolules obsolete
or distinct and to 15 mm; terminal leaflet subequaling lateral ones or larger and to 6.5 cm and to
7 cm wide, sometimes suborbicular or subreniform. Racemes ebracteate; fruiting racemes 3–
17(–25) cm. Fruiting pedicels (8–)10–18(–22) mm. Sepals (1.5–)3–4 mm, 1–1.5 mm wide;
petals white, obovate, (6–)8–10 mm, 4–6.5 mm wide; filaments of median stamens 5–6.5 mm,
those of lateral pair 4–5 mm; anthers 1.4–1.8 mm; ovules 12–16 per ovary. Fruits linear,
(1.3–)1.5–3.2(–4) cm, 1.4–2 mm wide, attenuate at apex; style 1.5–2(–2.5) mm. Seeds (1.2–)1.6–
2.2 mm, 1.2–1.6 mm wide, ovate, light brown. 2n = 16, 32, 46–48, 56, 72.
Flowering: May–Aug.
Habitat: wet meadows or open areas along mountain streams.
Elevation: 200–2000 m.
Distribution: China (Jilin), Japan (Hokkaido, N Honshu, Kyushu), Russia (Far East).
Cardamine yunnanensis Franchet, Bull. Soc. Bot. France 33: 398. 1886. TYPE: China,
Yunnan, Ta-long-tan, near Tapin-tze, 1800 m, 26 Jul 1885, J. M. Delavay 1843 (holotype, P!;
isotypes, E!, F!, P!, US!).
Cardamine bijiangensis W. T. Wang, Acta Bot. Yunnan. 9: 12. 1987. TYPE: China, Yunnan,
Bijiang, Kungdung, 2800 m, 9 Jul 1978, Nujiang Expedition 943 (holotype, KUN!; isotype,
KUN!).
Cardamine hirsuta var. oxycarpa J. D. Hooker & T. Anderson,
Cardamine heterophylla T. Y. Cheo & R. C. Fang, Bull. Bot. Lab. North-East. Forest. Inst.,
Harbin 1980(6): 27. 1980; not Host, Syn. P. Austral. 366. 1797; not Lapeyrouse, Hist. Abr. Pl.
Pyr. 377. 1813; not Bory, Ann. Sci. Gen. Phys. 3: 6. 1820; not W. J. Hooker, Comp. Bot. Mag. 1:
273. 1835; not (Nuttall) Wood, Amer. Bot. & Fl. 38. 1870; not (Forester) O. E. Schulz, Bot.
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Jahrb. Syst. 32: 487. 1903; Cardamine sinica Rashid & H. Ohba, J. Jap. Bot. 68: 182. 1993.
TYPE: China, Sichuan, Nanchuan Xian, Daheba, 13 Apr 1938, K. L. Chu 6034 (holotype,
NAS!).
Cardamine inayatii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 9: 1069. 1927. TYPE: India,
Sikkim, below Chiyabanjan, 2500 m, 11 May 1960, Hara et al. s.n. (holotype, TI!).
Cardamine levicaulis W. T. Wang, Acta Bot. Yunnan. 9: 9. 1987. TYPE: China, Sichuan,
Luding, Mons Gonggashan, Yanzigou, 2500−3000 m, 25 Jun 1982, Lang Kaiyong, Li
Liangqian & Fei Yong 226 (holotype, PE!; isotype, KUN!, PE!).
Cardamine longipedicellata Z. M. Tan & G. H. Chen, Acta Phytotax. Sin. 34: 650. 1996, not
Rollins, J. Arnold Arb. 21: 395. 1940. TYPE: China, Sichuan, Shifang, Saiyunhua, 1090 m, 3
Apr 1979, S. Huang 40 (holotype, SZ!).
Cardamine longistyla W. T. Wang, Acta Bot. Yunnan. 9: 14. 1987. China, Xizang, Zayü,
Ridong, Shiladu, 3600 m, 8 Sep 1982, Qinghai-Xizang Team 10183 (holotype, PE!; isotype,
KUN!).
Cardamine muliensis W. T. Wang, Acta Bot. Yunnan. 9: 11. 1987. TYPE: China, Sichuan, Muli,
Bailianggou, 3300 m, 26 Aug 1983, Qinghai-Xizang Team 13497 (holotype, KUN!).
Cardamine sikkimensis H. Hara, J. Jap. Bot. 37: 97. 1962. TYPE:
Cardamine weixiensis W. T. Wang, Acta Bot. Yunnan. 9: 9. 1987. TYPE: China, Yunnan,
Weixi, Lidiping, 3100−3300 m, 21 Jul 1981, Expedition of Hengduanshan 1828 (holotype,
PE!; isotype, PE!).
Herbs, short-lived perennial, rarely annual. Rhizomes slender. Stems (1–)1.5–4.5(–6) dm,
often pilose or puberulent, sometimes glabrescent distally, simple or branched from base, angled.
Basal leaves petiolate, often withered by flowering, 3–5-foliolate, rarely simple; petiole 1–6(–8)
cm; leaf blade or terminal leaflet suborbicular, ovate, or lanceolate, 0.5–3 × 0.5–2.5 cm, dentate,
crenate, or rarely subsinuate; lateral leaflets absent or 1 or 2 pairs. Middle cauline leaves 3–7foliolate; petiole (1–)2–7(–8.5) cm, basally auriculate; auricles toothlike, linear, or lanceolate,
(0.4–)1–3(–4) × 0.2–1(–1.5) mm; terminal leaflet lanceolate, elliptic, oblong, ovate, or
suborbicular, (1–)1.3–4.5(–6) × (0.4–)0.6–2(–3) cm, sparsely pilose adaxially, often glabrous
abaxially, with a petiolule (2–)4–14(–20) mm, base obtuse or cuneate, margin ciliate to ciliolate
and dentate, crenate, sinuate, or rarely repand, apex acute or acuminate; lateral leaflets 1–3 pairs,
shortly petiolulate to subsessile, similar to terminal one and often oblique at base; uppermost
leaves often trifoliolate, rarely simple. Raceme ebracteate; fruiting pedicels ascending or
divaricate, 0.5–1.8(–2.3) cm, straight, slender. Sepals oblong or nearly ovate, 2–3 × 1–1.5 mm,
base not saccate; petals white, obovate, (2.5–)3.5–5(–6) × 2–3 mm, not clawed; median filament
pairs 2.5–4 mm, lateral pair 2–3 mm; anthers oblong, 0.5–0.8 mm; ovules 8–18 per ovary. Fruit
linear, 1.5–2.8(–3) cm × 1–1.3 mm; valves smooth, sparsely pilose; style (0.5–)1–2.5(–3.5) mm.
Seeds brown, narrowly oblong, 1.3–1.8 × 0.7–1 mm, wingless.
Flowering: Mar–Jul.
Habitat: moist shady places, mountain slopes, valleys, grasslands, meadows, thickets, forest
openings, damp stream beds.
Elevation: 900–4200 m.
Distribution: Bhutan, China (Sichuan, Xizang, Yunnan), India (Sikkim), Nepal.
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